Issue No. 16 Newsletter of the Pineapple Working Group, International Society for Horticultural Science
Table of Contents
July, 2009
New s of General Interest, Pineapple W orking Group New s. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
‘MD-2’ Pineapple Transforms the World’s Pineapple Fresh Fruit Export Industry. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
7th International Pineapple Symposium. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
ISHS Publishes Proceedings of International Pineapple Symposium VI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Contribute to Pineapple News. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
New s from Australia.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
New s From Belgium.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
A Novel Flowering Induction Agent for Pineapple. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
MD-2 Pineapple Plant Maturity and Flowering. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
New s From Brazil. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
New Records of Scale Insect Pests of Pineapple and Their Natural Enemies. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
New s From Cuba. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
Cryogenic Strategy For The Establishment Of Pineapple (Ananas Comosus L. Merrill) Germplasm Bank. . . . . . . . . . . . . . . 14
Phenotypic Characterization of Field-grown Pineapple Transgenic Plants. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
Proteases Expressed in Response to in Vitro Culture of Pineapple. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
New s From France. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
The Domestication of Pineapple: Context and Hypotheses. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
New s from the Philippines. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
Philippines President Witnesses Signing of $500-M Contract. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
New s from Sri Lanka. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
Antagonistic effect of Trichoderma harzianum on Thielaviopsis paradoxa. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
New s from Taiwan. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
Forced Flowering of Pineapple (Ananas comosus cv. Tainon 17) With Calcium Carbide Plus Activated Charcoal and By Ice
Cold Stress.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
New s From the United States (Haw aii).. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
Phytophthora Heart Rot Control. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
Evaluation of Transgenic Resistance. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
Production of transgenic pineapple (Ananas cosmos (L.) Merr.) plants via adventitious bud regeneration. . . . . . . . . . . . . . . 34
Effects of ReTain® on Natural Induction of Reproductive Development of MD-2 Pineapple. . . . . . . . . . . . . . . . . . . . . . . . . . . 34
Services.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
Commercial Services. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
Professional Services. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
Book Review s and W eb Sites. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
Book Reviews. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
Web Sites of Possible Interest. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
Instructions to Contributors to Pineapple New s.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
Newsletter of the Pineapple Working Group, International Society for Horticultural Science
News of General Interest, Pineapple Working Group News
Dear Colleagues:
Another year has gone by and we are now looking forward to hearing about plans for symposium VII. Content for the
newsletter is down this year, presumably in part due to the fact that it takes time to generate research results for pineapple and
because the number of researchers on the crop also continues to dwindle. Graham Petty a well-know pineapple researcher in
South Africa, retired this past year. Pineapple growers there are having a difficult time competing with low-cost producers in
equatorial Africa and it seems unlikely that his position will be filled in the near future. Despite such changes, this issue includes a
number articles that I hope readers will find interesting as well as a sizable list of recent references on pineapple and related
topics.
For quite a number of years I have been curious how the domination of ‘Smooth Cayenne’ in the fresh fruit pineapple market
was broken. Dr. Robert E. Paull and I nominated ‘MD-2' pineapple for the American Society for Horticulture Science 2009
Outstanding Fruit Cultivar Award. W hile collecting information for that nomination I also was able to obtain additional
information about the history leading to the development of ‘MD-2’. Below is the story of the transformation based on the best
information I have been able to collect.
‘MD-2’ Pineapple Transforms the World’s Pineapple Fresh Fruit Export Industry
Duane P. Bartholomew. Dept. Tropical Plant & Soil Sci., Univ. Hawaii at Mano`a, Honolulu, HI 96822. E-mail:
duaneb@hawaii.edu.
The world’s pineapple fresh fruit export industry went through a remarkable transformation after Del Monte Corporation
introduced ‘MD-2’pineapple to consumers in the United States and Europe, officially in 1996 (Frank, 2003). Until the mid 1990s,
the world’s fresh fruit export industry was relatively small and was based mainly on cultivar Smooth Cayenne. Prior to the
introduction of ‘MD-2’, the focus of the pineapple export industry was on canned ‘Smooth Cayenne’ pineapple. W ith the major
producers focused on ‘Smooth Cayenne’ pineapple, the unanswered question is what circumstances made it possible for Del
Monte Corporation to became the first multinational pineapple company in the world to break the industry fixation on ‘Smooth
Cayenne’ and risk growing a new and untested hybrid pineapple?
Some Industry History
As the world’s economies recovered after W orld W ar II, the pineapple growers in Hawaii faced increasing competition from
low-cost producers in the Philippines and Thailand. Del Monte Corporation had established a pineapple plantation and cannery in
the Philippines before W orld W ar II but the Hawaii business remained mostly profitable until the late 1970s. As prices for canned
pineapple dropped due to increased foreign competition in the 1950s and 1960s, the Hawaii plantations began to close. By 1980,
only Del Monte, Dole and Maui Pineapple Company remained in the canning business in Hawaii. Hawaii’s glory days as the
world’s leading canner of pineapples was nearing its end. Del Monte, the only large company in Hawaii that farmed on leased
land, closed their Hawaii cannery in 1983 and shifted all canning operations to the Philippines and Kenya. Dole also scaled down
their Hawaii operation and eventually closed their cannery in 1991. W ith the closure of the Del Monte cannery, the company’s
options were to close the plantation or concentrate on growing and marketing fresh fruit, a major transformation because it
involved developing a year-around production system that was based on a mostly full-time labor force.
Production of fruit for the cannery was concentrated in the summer months because fruit quality was better and seasonal
labor, primarily students out of school, was more readily available to man the harvesting and canning operations. W hen Del
Monte moved to an all fresh fruit production system, one of the issues the company was forced to confront was the seasonal
variation in fruit quality. It had long been known that pineapple fruit quality was lower in the winter because fruit acidity was
much higher during that time of year. But the major holidays that fell during the winter occurred at times when the variety and
supply of other fresh fruits was reduced and thus were viewed as important market opportunities. If the quality of ‘Smooth
Cayenne’ fruit had been consistent throughout the year, consumers would have been unable to compare high quality summer fruits
with the lower-quality fruit produced in the winter. Thus it is likely that Del Monte management knew that something superior to
‘Smooth Cayenne’ was needed if their pineapples were going to be competitive with other fresh fruits for the consumers dollars.
W here was such a superior cultivar to come from?
In 1961, the Pineapple Research Institute of Hawaii (PRI) was asked to extend its purview beyond field operations to explore
uses for pineapple that might expand the market for the fruit (Gortner et al., 1963). One of the “new products” thought to have
potential was field-ripe pineapple for the fresh fruit markets of the United States. Fresh fruit was first shipped from Hawaii to U.S.
markets in 1849 and fruit continued to be shipped from Hawaii over the years. However, the focus of the industry in the early 20 th
century and onward was on processed pineapple. As a result of many years of experience with ‘Smooth Cayenne’, the mind set of
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Newsletter of the Pineapple Working Group, International Society for Horticultural Science
company managements was that any fruit shipped from Hawaii must look and taste like ‘Smooth Cayenne’ to be acceptable.
There is more than one account of a plantation or company manager rejecting what was thought to be a promising new hybrid
because it didn’t look or taste like ‘Smooth Cayenne’. In a discussion of how Hawaii might compete for the fresh fruit market
with countries in the Caribbean and Central America, the Gortner et al. (1963) report stated “Hawaii has exclusive and distinct
varieties available to choose from, and may be able to offer a variety better adapted to the fresh fruit market demands.” Despite
that claim, the focus of the PRI research report was on ‘Smooth Cayenne’ and the focus of Hawaii’s pineapple producers remained
on ‘Smooth Cayenne’ for another 20 years. No new pineapple cultivar well suited to the fresh market was being grown in more
than small-scale tests.
It is understandable that a company that was mainly in the business of growing pineapple for canning would be reluctant to
explore new cultivars. ‘Smooth Cayenne’ pineapple was highly productive, was reasonably resistant to the important pests and
diseases, and produced a fruit that met consumer expectations when canned. Changing a large plantation over to a new cultivar is
very costly and takes many years. A gradual changeover of cultivars also poses significant management problems. Care needs to
be taken to prevent mixing of planting material in the field and of mixing fruits in the cannery, especially if the fruit of the new
cultivar has a different appearance in the can. Growing different cultivars on the same farm also increases the chances of natural
crossing, which would produce fruits with seeds. In a quality-conscious industry, seedy fruits would be rejected in the cannery,
resulting in significant losses and increased costs. Further, Hawaii growers were by nature cautious because catastrophic failures
were the common experience. In the early 1960s, PRI hybrid 53-116 appeared to be so superior to ‘Smooth Cayenne’ that the
main Hawaii plantations had planted several hundred acres of that hybrid. Despite more than 10 years of testing, a fatal flaw,
susceptibility to a fruit disease, was exposed that resulted in large losses of fruit in the cannery. The hybrid was instantly
discarded and no doubt industry skepticism about the possibility of replacing ‘Smooth Cayenne’ with a more productive or higher
quality hybrid was heightened.
W hen W illiams and Fleisch (1993) presented a historical review of pineapple breeding programs in Hawaii, including that of
the PRI, they gave no indication that a hybrid from the 58 year pineapple breeding program would produce a cultivar that would
transform the world’s pineapple fresh fruit export industry. There is no indication from their paper that they had any knowledge
that the transformation process had already begun on a new plantation established by Del Monte Corporation in Costa Rica. ‘Even
as the PRI breeding and selection program ended in the mid 1980s, the focus of the program was on the breeding and selection of
cultivars that would can well rather than on fresh fruit cultivars. It was up to George Yamane, Research Manager at Del Monte
and, after his death, Calvin Oda, to see the transformation process through to its astoundingly successful conclusion.
The Story of ‘M D-2’
W illiams and Fleisch (1993) reported that PRI hybrids had been developed that, when compared with ‘Smooth Cayenne’ had:
•
better resistance to Phytophthora rots, pineapple wilt, nematodes, pink disease and internal brown spot
•
higher levels of vitamins C and A
•
reduced acid content in winter ripened fruit
•
better harvest peaking
•
higher yield
•
better cannery recovery from a ton of fruit
•
faster plant growth
•
a range of distinctive flavors.
During intensive screening, all of these varieties exhibited flaws that prevented them from replacing ‘Smooth Cayenne’, either
for canning or for fresh fruit. After 50,000+ seedlings screened from each of the most promising parental crosses made in the late
1960's - early 70's failed to produce a commercially acceptable cultivar, the decision was made to phase out the PRI breeding
program. Crossing was terminated in 1972, the PRI experiment station was closed in 1975 and evaluation of all seedlings was
completed at Maui Pineapple Company in 1985. Several PRI selections were released to PRI member companies for potential
commercial use.
Dr. David D.F. W illiams and his able assistants, Frank Bermudas and Toshio Minagawa, made many crosses in 1970 but one
cross was destined to make history. That was a cross between the PRI hybrids 58-1184 and 59-443, both of which had superior
characteristics but had problems that prevented them from becoming commercial successes. The seeds of that cross were planted
in 1971 and among the plants that grew to produce fruit, at least two were selected as having promise in 1973. During further
evaluation, those two hybrids were carried in the breeder's book as selections 50 and 114, i.e. 73-50 and 73-114. The parentage of
the parents of the two clones are relatively complex as pineapple parents go, being mixtures of ‘Smooth Cayenne', Smooth
Guatemalan, ‘Ruby’ (a ‘Spanish’ clone), ‘Queen’ and ‘Pernambuco’ (W illiams and Fleisch, 1993). Both are more than 50%
‘Smooth Cayenne', which is significant for Hawaii and the U.S. mainland because they are not considered potential fruit fly hosts
(Armstrong et al., 1979; Seo et al., 1973). Between 1973 and 1980 test plantings of the two hybrids, perhaps along with others
that showed some potential, were made on Maui and, between 1978-80, also on the Del Monte plantation on Oahu.
The hybrids were released by the PRI to the then member companies Del Monte Corp. and Maui Pineapple Co. for further
evaluation in 1980. In 1981 Del Monte renamed 73-50 as “MD-1", though it was never known by that name outside of Del
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Newsletter of the Pineapple Working Group, International Society for Horticultural Science
Monte, and 73-114 was named “MD-2". Both were named after Del Monte Hawaii general manager Frank Dillard’s wife Millie
by Del Monte’s research manager George M. Yamane and his assistant Calvin Oda. Hybrid 73-50, which due to a patent that was
obtained for what has been assumed to be that hybrid, is perhaps now most widely known as ‘CO-2’ (Chan et al., 2003). ‘CO-2’
has achieved some limited success, but no where near that enjoyed by ‘MD-2’. ‘MD-2’ is a vigorous plant with acceptable disease
and pest resistance, high fruit yield and outstanding fruit quality. The outstanding fruit characteristics include golden external
shell color, golden flesh color, slightly higher soluble solids content than ‘Smooth Cayenne', considerably lower acidity, especially
during the winter months, higher vitamin C content, and exceptional post-harvest shelf life. No other pineapple cultivar in the
international pineapple trade holds up as well under refrigeration as does ‘MD-2’. These attributes of ‘MD-2’ apparently
convinced Yamane and Oda to support expanding plantings of the hybrid. In addition, containers of planting material were
shipped to Del Monte’s Philippines subsidiary in 1983-84. Test marketing of ‘MD-2’ was done in Japan, but acceptance there was
slow because the cultivar was very different from ‘Smooth Cayenne’ and had not yet become popular elsewhere.
As part of Del Monte’s plan to expand their fresh fruit business, the company took over an abandoned pineapple farm of
several hundred hectares in Costa Rica in the late 1970s. Armed with the knowledge that select clones of ‘Smooth Cayenne’ are
superior to field-run plants, Del Monte replaced the local clone with plants of the PRI ‘Smooth Cayenne’ clone Champaka F-153,
which at the time was being grown in Hawaii by both Del Monte and Maui Pineapple Company. The new clone performed
poorly, which is not an unusual finding for a clone selected for the Hawaii environment. Champaka F-180, also part of the PRI
‘Smooth Cayenne’ clone collection, was inferior to Champaka F-153 in Hawaii. However, F-180 was superior the F-153 clone in
the colder environment of Queensland, Australia and became an important clone there.
In an attempt to improve production on their new plantation, company researchers Yamane and Oda had plants of ‘MD-2’
shipped from Hawaii to Costa Rica in the mid 1980s. The details of how development of ‘MD-2’ progressed in Costa Rica are not
publicly available; however, Frank (2003) speculated that test marketing was done in the late 1980s or early 1990s when fruits
were sold by what was then Del Monte Tropical Fruit (Fresh Del Monte Produce, 2009) in Boston and Florida under the label Del
Monte Gold® Extra Sweet (for a time the boxes also carried the ‘MD-2’ label (D. Bartholomew, personal observation)). It is
likely that as a result of these early test marketings, Del Monte management realized that the company had a winner on its hands.
Sale of ‘MD-2’ fruits in Europe followed not long after it was introduced in the U.S. market.
The period from about 1993 until 2003 is surrounded with intrigue (Frank, 2003; Janick, 2003; Greig, 2004) that has little to
do with the success of ‘MD-2’ and only impacts which companies were able to benefit from it, a not insignificant issue. W hat is
now widely known is that fruits of ‘MD-2’ were so well accepted that they were sold at premium prices for about 10 years and, in
that same time frame almost completely displaced ‘Smooth Cayenne’ as the world’s main fresh fruit cultivar grown for export (see
details below).
During the period when Fresh Del Monte Produce (the name was changed in 1996) was the sole source of ‘MD-2’, pineapple
sales reported by the company increased from nearly $200 million dollars in 1996 to $440 million in 2002, the last year the
company broke out pineapple sales in their annual report (Frank, 2003). ‘MD-2’ is now grown by many companies and growers,
both large and small, is the worlds’ main fresh fruit cultivar and the fruit is exported to temperate markets in the United States,
Europe, England, Japan, Korea, Hong Kong, China, Singapore and the Middle East.
Statistics on the area planted to ‘MD-2’ and its market value are difficult to obtain because it is grown or marketed, or both,
primarily by multinational corporations. The area planted to ‘MD-2’ exceeds 60,000 hectares (ha) (FAO Ag Stat 2007 and other
sources). Costa Rica alone has 42,500 ha and is the country with the largest area planted to ‘MD-2’. Other countries and
estimates of areas planted to ‘MD-2’ include Ecuador (5,850 ha), Honduras (2,800 ha), Guatemala (500 ha), Panama (1,600 ha),
Brazil (>500 ha), Ghana (>600 ha), Cote d'Ivoire (>500 ha), Hawaii (about 820 ha) and Philippines (approximately 10,000 ha in
the Davao and Bukidnon areas; the report is that one large grower plans to plant several thousand more hectares in the next five
years).
Both the production and consumption of ‘MD-2’ pineapple continue to grow. Loeillet (2008) reported that in 2006 Costa
Rica alone exported nearly 1.2 million metric tons (MT) of fresh pineapple, a 30% increase over 2005; increases in shipments
were +25% from 2003 to 2004 and +30% from 2004 to 2005. Exports from Costa Rica have multiplied by 2.6 since 2002 and it is
unlikely that such almost explosive growth in fresh sales have ever been recorded for any other fruit or fruit cultivar.
In addition to the rapid growth of ‘MD-2’ in the marketplace, its success had a devastating impact on pineapple sales from
Cote d’Ivoire (Loeillet, 2008) and Ghana. In about the year 2000, Ghana experienced a dramatic drop in pineapple exports
because the market favored ‘MD-2’ rather than ‘Smooth Cayenne', Ghana's principle export cultivar. In 2002, with $2 million of
government support, tissue culture production of ‘MD-2’ was rapidly ramped up and by the end of 2007 Ghana had exported
42,000 MT of ‘MD-2’, earning $20 million in foreign currency. As a result, Ghana was ranked 3 rd as an exporter to the EU
(Ghana Export Promotion Council, 2008).
Based on the information obtained for this paper, it would appear that there is no simple answer to the question posed at the
beginning of the article. ‘MD-2’ pineapple was introduced at a time when consumer preferences were changing, which resulted in
a rapid increase in the consumption of fresh fruits and vegetables. If Del Monte had found success with ‘Smooth Cayenne’ in
Costa Rica, perhaps another company would have been the first to introduce a new pineapple cultivar to the world’s consumers.
But all credit must go the Del Monte researchers Yamane and Oda, or the company’s managers, or both, because they were quick
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Newsletter of the Pineapple Working Group, International Society for Horticultural Science
to recognize and capitalize on ‘MD-2’ once it became clear that consumers preferred it over any other pineapple cultivar then
offered in the United States or Europe. Once ‘MD-2’ was accepted in the U.S. and EU markets, acceptance was rapid in Asian
markets too. As a result of these remarkable successes, ‘MD-2’ is the first major pineapple cultivar to be produced by modern
breeding. It has now essentially replaced the ancient ‘Smooth Cayenne' developed by Amerindians in pre-Columbian times as the
world's principal pineapple fresh fruit cultivar grown for export. Though the total market value of ‘MD-2' fruits is not known, it is
likely that annual sales of that cultivar alone exceed $1 billion.
Dr. David D. F. W illiams is now comfortably retired in the state of Colorado. In July of 2009 W illiams will travel to the
American Society for Horticulture Science (ASHS) annual meetings in St. Louis, Missouri to receive a medal recognizing the
contributions of him and his predecessors at the Pineapple Research Institute of Hawaii in the development of PRI hybrid 73-114.
And this hybrid, now known widely as cultivar ‘MD-2’, will be recognized as the ASHS 2009 Outstanding Fruit Cultivar. This
article is based in part on the nomination of ‘MD-2’ for that award, which Duane P. Bartholomew and Robert E. Paull had the
pleasure of preparing. W ith the transformation begun by ‘MD-2’, perhaps at some future ASHS or other horticultural society
meeting, another new pineapple cultivar will receive an award as the outstanding cultivar and consumers will have an opportunity
to choose from several pineapple cultivars produced by modern breeding.
References
Armstrong J.W., Vriesenga J.D., Lee C.Y.L. (1979) Resistance of pineapple varieties d-10 and d-20 to field populations of oriental fruit flies
and melon flies. J Econ Entomol 72:6-7.
Chan Y.K., Coppens d'Eeckenbrugge G., Sanewski G.M. (2003) Breeding and variety improvement 33-55, in: D. P. Bartholomew, R.E. Paull
and K.G. Rohrbach (Eds.), The Pineapple: Botany, Production and Uses, CABI Publishing, Wallingford. pp. 320.
Frank, R. 2003. Going for 'The Gold' Turns Pineapple World Upside Down. Wall Street Journal, Oct. 7, p. 1.
Fresh Del Monte Produce. 2009. http://www.freshdelmonte.com/ourcompany/companyoverview/history.aspx. Accessed on July 4, 2009.
Loeillet, D. 2008 (March). Close-up – FruiTrop. (http://passionfruit.cirad.fr/index.php/recherche/(produit)/10) No. 154. 30 pages.
Ghana Export Promotion Council. 2008. http://www.gepcghana.com/news.php?news=21. Accessed July 4, 2009.
Greig, I. 2004. Pineapple Wars Redux. Chronica Horticulturae 44 (2), 5
Gortner, W.A., Spiegelberg, C.H., Dull, G.G., and Krauss, B.H., 1963. Field-fresh pineapple for export. Research Report 99.
Janick, J. 2003. Pineapple Wars. Chronica Horticulurae 43 (4), 17
Seo S.T., Chambers D.L., Lee C.Y.L., Komura M., Fujimoto M., Kamakahi D. (1973) Resistance of pineapple variety 59-443 to field
populations of oriental fruit flies and melon flies. J Econ Entomol 66:522-523.
Williams, D.D.F. and Fleisch, H., 1993. Historical review of pineapple breeding in Hawaii. Acta Horticulturae 334:67-76.
7th International Pineapple Symposium
At the Pineapple W orking Group meeting at the VIth Symposium it was agreed that the VIIth International Pineapple
Symposium would be held in 2010. The symposium is to be organized by the Malaysian Pineapple Industry Board (MPIB) and
the Malaysian Agricultural Research and Development Institute (MARDI) with the support of the Ministry of Agriculture and
Agro-based Industry Malaysia (MOA). Mr. Tengku Malik (tamtm@mardi.gov.my) will lead the organization of this symposium.
No formal announcement has come from MARDI to date (mid May, 2009). Malaysia is an interesting country to visit and offers
many amenities as well as beautiful and unique local crafts.
ISHS Publishes Proceedings of International Pineapple Symposium VI
The proceedings of the VIth symposium were recently published and the volume can be obtained from the International
Society for Horticultural Science web site at http://www.ishs.org. Plan to visit the much updated web site soon. The ISHS is one
of the foremost organizations promoting cooperation and communication among horticultural researchers, growers and consumers.
The ISHS continues to expand its offerings to members as well as to provide the structure under which our Pineapple W orking
Group (http://www.ishs.org/science/T07.php) functions. Detailed information about ISHS and the benefits of membership can be
found at http://www.ishs.org or you can write to the ISHS Secretariat, P.O. Box 500, 3001 Leuven, Belgium (E-Mail:
info@ishs.org).
Contribute to Pineapple News
Information on how to contribute to Pineapple News can be found at the end of the newsletter. You can also contact Duane
Bartholomew, the editor, at duaneb@hawaii.edu.—
News from Australia
No articles were submitted from Australia but Annual Industry Reports covering work on pineapple in the country for the
years 2006-07 and 2007-08 can be obtained at http://www.horticulture.com.au/industry/annualreports.asp.
5
Newsletter of the Pineapple Working Group, International Society for Horticultural Science
Topics covered in the 2006-07Report include:
•
Industry Tackles Challenges Through R&D
•
Queensland Industry Manages Changes
•
Pineapple Industry Biosecurity Plan Update
•
Levies to Develop the Australian Pineapple Industry
•
Across Industry Program 2006/07
•
Pineapple Program 2006/07
Topics covered in the 2007-08 Report include:
•
Industry supports new national levies
•
Navigating change in the Queensland pineapple industry
•
Development of new local fresh market pineapple cultivars
•
Establishment of levies to strengthen the Australian pineapple industry
•
Generation of dimethoate residue data to support ongoing use in pineapple
•
Pineapple study tour of Brazil and Costa Rica
•
Across industry program 2007/08
•
Pineapple program 2007/08
•
Investing in a Astralian horticulture
—
News From Belgium
A Novel Flowering Induction Agent for Pineapple
Van de Poel B.1 *, Ceusters J.2, De Proft M.P.2
1
BIOSYST-MeBioS, Department of Biosystems, Faculty of Bioscience Engineering, Katholieke Universiteit Leuven, W illem de
Croylaan 42, B-3001 Leuven, Belgium
2
Division of Crop Biotechnics, Department of Biosystems, Faculty of Bioscience Engineering, Katholieke Universiteit Leuven,
W illem De Croylaan 42, B-3001 Leuven, Belgium. *Corresponding author E-mail: bram.vandepoel@biw.kuleuven.be
Abstract
This paper is condensed from a recently published article "Determination of pineapple (Ananas comosus, MD-2 hybrid
cultivar) maturity, the efficiency of flowering induction agents and the use of activated carbon" published in Scientia Horticulturae
120 (1) page 58-63. W e report on a novel flowering induction agent called "zeothene", which releases gaseous ethylene on contact
with water. W e evaluated the induction capacity of this agent on a commercial plantation (Ecuador) and compared its flowering
induction efficiency with other common agents currently used in pineapple cultivation. The zeothene treatments resulted in
homogeneous flowering, which is highly desired on commercial farms. Furthermore we evaluated the method of induction (central
cup application vs. whole plant spraying) and the use of activated carbon to enhance the flowering induction treatment with
ethylene gas. Our results indicated that a central cup application is more favorable to obtain a homogeneous flowering and that the
beneficial effects of activated carbon are questionable.
Introduction
Pineapple (Ananas comosus) is a member of the Bromeliaceae family and is characterized by its ability to flower in response
to ethylene signals (internal or external). This feature is exploited worldwide by pineapple growers to synchronize flower and fruit
development, which reduces harvesting costs and optimizes market supply (Min & Bartholomew, 1996). It is very important to
apply the correct flowering induction agent at the right time. This is a crucial aspect in pineapple cultivation to maximize the
number of plants fruiting at the same time and, consequently, to maximize yield. In this report we focus on the correct use of
different flowering induction agents.
Ethephon (2-chloroethylphosphonic acid) is a common flowering induction agent. The chemical is stable under acidic conditions
(pH #3) and breaks down to release ethylene gas at a higher pH ($5) (Yang, 1969; Dass et al., 1976). Usually it is applied as a
broadcast spray with a boom sprayer during the day or, if temperatures are high, in the late evening or at night to prevent
evaporation losses (Turnbull et al., 1999). Increased amounts or multiple applications are used if plants are expected to have low
sensitivity.
6
Newsletter of the Pineapple Working Group, International Society for Horticultural Science
Another very common flowering induction agent is ethylene gas. Ethylene has been proven more efficient than ethephon (Py
et al., 1984). The gas is injected into water under pressure and applied as a broadcast spray over the plants with a boom sprayer.
Activated carbon is added to the water to increase the volume of gas retained in the solution. Ethylene gas is most effective when
applied late in the evening, at night, or in the early morning. Such timing reduces evaporation losses and the lower temperature at
night increases the solubility of the gas in the water droplets (L'Air Liquide, 1976). It is also believed that ethylene uptake by the
plant is higher at night because during this period the stomata are open (due to the Crassulacean Acid Metabolism of the plant)
(Bartholomew et al., 2003). A suitable ethylene concentration used for commercial flowering induction is 2.272 kg ha -1 ethylene
gas sprayed with 7000 L ha-1 water containing 20 kg activated carbon/ha (Hepton, 2003).
The aim of this research was to evaluate the flowering induction efficiency of a novel agent, called "zeothene". This was done
on a commercial plantation. Application was directly into the central cup with and without activated carbon.
M aterials and methods
Flowering induction experiment
All field experiments were executed on a commercial pineapple plantation near Quevedo (Ecuador). The plantation was a
monoculture of the new hybrid cultivar MD-2. The efficiency of four different flowering induction agents was tested (zeothene,
ethephon, ethylene gas and ethylene dissolved in water). All treatments were applied in the early evening (5-6 p.m.) to plants of
8-9 months after planting. "Zeothene" is a special zeolite pearl retaining ethylene gas, which is released upon contact with water.
This zeothene (or also called 'ethylene pills') was developed by M.P. De Proft (Laboratory of Plant Production, Department of
Biosystems, Katholieke Universiteit Leuven, Belgium). The pills (± 13.5 mg/pill containing ± 0.7 mL C 2H 4) are very effective in
inducing flowering in ornamental bromeliads (Parton, 2001; Plever, 2004). To induce flowering of pineapple, 3-4 pills were
dropped into the central cup of each plant. The pills release their ethylene content upon coming in contact with water standing in
the central cup of the plants.
Ethephon (Ethrel, Bayer CropScience, Germany) was provided with a commercial dose of 0.5 kg ethephon per hectare (active
ingredient) dissolved in 2000 L water with 5 % urea and brought to pH 3. A small amount of the prepared ethephon solution was
brought into the central cup with a hand sprayer. A solution containing the equivalent of 2.272 kg ethylene gas in 7,000 L of water
with 20 kg activated carbon (Hepton, 2003) was applied over the entire plants by a pressurized boom sprayer. This is the standard
flower induction application of the plantation. The ethylene-water solutions were made by adding a known amount of zeothene
pills to a sealed 25 L tank containing water, resulting in the following concentrations of ethylene, C 2H 4 in g L -1: 0.389, 0.292 and
0.195, corresponding respectively to 100%, 75% and 50% of the commercial dose used for ethylene gas. Activated carbon (2.86 g
L-1 ) was added to two of the three doses tested. The prepared solutions were applied into the central cup of the plant with a hand
sprayer. All treatments were applied randomly to rows of 100 plants and the experiment was repeated 3 times.
The efficiency of the flowering induction treatment can be expressed by a flowering homogeneity percentage. This percentage
was obtained by counting the number of plants which were in the same floral development stage exactly 73 days after induction.
W e distinguished 6 main floral stages: opening of the central cup, red bud stage (= inflorescence tip is visible), 1st flower (=
lowest petals are visible), 2nd flower (= middle petals are visible), 3rd flower (= upper petals are visible) and the wilting stage (=
all petals are wilted) (Rohrbach & Taniguchi, 1984).
Ethylene absorption by activated carbon
Ethylene absorption of different concentrations of activated carbon-water solutions (5%, 0.5% and 0.05%; Pro-analyse, Vel
NV, Belgium) was tested by means of gas chromatography. One hundred mL of each solution was added to a 300 mL airtight
glass flask and a known amount of ethylene gas (1.25 ppm) was injected in the headspace of the flask. The solutions were stirred
gently (150 RPM) and air samples were taken at regular time intervals. The air samples were analyzed for their ethylene content
by a gas chromatograph (DI 200, Delsi Instruments, France) equipped with a flame ionization detector (FID) (De Greef & De
Proft, 1978). Each concentration was tested 3 times. All measurements were conducted at normal room conditions (23°C and 1013
hPa).
Results
Flowering induction experiment
Homogeneity of induction results (Table 1) show that "zeothene" successfully induced flowering though it was only
significantly better than the ethylene gas treatment (Table 1). W hile other treatments were not different from each other, flowering
stage development of plants forced with ethephon (Table 1) was delayed compared with the other treatments. This might be
explained by the fact that ethephon first has to break down into ethylene before it can exert its physiological action. The
commercial ethylene application had the lowest homogeneity percentage of all treatments. This low percentage will eventually
result in a more heterogeneous flower and fruit development which consequently may result in multiple harvest passes.
7
Newsletter of the Pineapple Working Group, International Society for Horticultural Science
A water solution containing dissolved ethylene gas with and without activated carbon also induced flowering in pineapple
when sprayed in the central cup. There was no significant effect of ethylene concentration and the addition of activated carbon to
the ethylene solution also had no significant effect on the homogeneity percentage. These results do not provide sufficient
evidence about the influence of activated carbon on the flowering induction reaction. Further research in this area is needed.
Ethylene absorption by activated carbon
To further investigate the role of activated carbon, the amount of ethylene absorbed by different activated carbon solutions
was quantified. Ethylene absorption isotherms (Figure 1) for three concentrations of activated carbon showed there was very
strong absorption of ethylene during the first 5 minutes and an equilibrium was reached after about 30 minutes of exposure. After
60 minutes the 5% activated carbon solution absorbed 65 % more ethylene gas than pure water (0.043 µl C2H4/100 ml compared
to 0.026 µl C2H4/100 ml) and the absorption was significantly greater after an exposure time of at least 20 minutes. Solutions
containing 0.5 or 0.05% activated carbon did not absorb significantly more ethylene gas than water itself, during the entire
exposure period.
Discussion and Conclusions
The effectiveness of flower induction mainly depends on the type of flower induction agent, the mode of application, plant
genotype and environmental conditions (temperature, humidity, wind, rain… ) (Bartholomew et al., 2003). The novel flowering
induction agent "zeothene" proved to be very efficient in inducing flowering in pineapple. Zeothene had a higher homogeneity
percentage than the ethephon and ethylene gas treatments. A solution of water with dissolved ethylene gas sprayed in the plant cup
resulted in a higher homogeneity percentage than did the “commercial” broadcast spray of ethylene in water with activated carbon
treatment; however, the higher percentage could be due to the method of delivery, i.e. into the central cup where as the
commercial treatment was a broadcast spray.
In these experiments we only investigated flowering homogeneity. W e did not take into account eventual fruit weight and
quality. These aspects should also be evaluated in future research before any final conclusions can be made about the new agent.
Nevertheless, it can already be stated that zeothene is a promising alternative to the currently used flowering induction agents.
Our field experiments clearly showed that central cup applications increase the homogeneity percentage and Turnbull et al.
(1999) found similar results for ethephon treatments. Central cup applications have the advantage that the active ingredient is near
or at the active site and is more protected from sun and wind so evaporation losses are smaller than for broadcast spraying. It is
the physiologically-active apical tissue that will have to change from a vegetative into a generative status. So ethylene perception
at the level of the apical meristem is a crucial aspect in pineapple flowering.
It is assumed that activated carbon enhances the flowering induction treatment by absorbing more ethylene gas, and thus
increasing the total amount of ethylene gas that is delivered to the plant. Therefore this is a common practice in pineapple
cultivation, although no published data on the subject could be found. Based on our field experiments no clear conclusions could
be made about the influence of activated carbon on the flowering induction treatment. Recently preliminary field trials conducted
by Lin (2008) have shown similar benefits of activated carbon in forcing. From our second experiment we can conclude that only
very high (5%) concentrations of activated carbon absorb significantly more ethylene than does pure water under laboratory
conditions without any applied pressure. During a commercial field application (0.286% activated carbon) where ethylene is
sprayed with water, the contact time between the gas and the water droplets is only limited, although a lot of pressure is applied. In
this short time frame it is questionable if any increased ethylene absorption is possible by the added activated carbon. Further
research on the influence of activated carbon is desirable. Field trials testing different types of commercial equipment and methods
of spraying can help illuminate this intriguing phenomenon.
Acknowledgments
The authors wish to thank José Ruiz and Luiz Fernando Saltarén for their expertise and guidance. This research was partly
supported by a grand from the Vlaamse Interuniversitaire Raad and the Katholieke Universiteit Leuven.
References
Bartholomew, D.P., Paull, R.E., Rohrbach, K.G. (Eds.) The pineapple: botany, production and uses. CAB International, Wallingford, pp. 320.
De Greef, J.A., De Proft,M., 1978. Kinetic measurements of small ethylene changes in
an open system designed for plant physiological studies. Physiol. Plant. 42, 79-84.
Dass, H.C., Randhawa, G.S., Singh, H.P., Ganapathy, K.M., 1976. Effect of pH and urea on the efficacy of ethephon for induction of flowering
in pineapple. Scientia Hort. 5, 265-268.
Hepton, A., 2003. Cultural systems. In: Bartholomew, D.P., Paull, R.E., Rohrbach, K.G. (Eds.), The pineapple: botany, production and uses.
CAB International, Wallingford, pp. 109-142.
L'Air Liquide, 1976. Encyclopedie des gaz. Elsevier, Amsterdam.
Lin, C.H., S. Maruthasalam, L.Y. Shiu, W.C. Lien, M. Loganathan, C.W. Yu, S.H. Hung, Y. Ko and Y.Y. Chen. 2008. Physical and chemical
manipulation of flowering in pineapple. Acta Horticulturae, 822, 117-123.
8
Newsletter of the Pineapple Working Group, International Society for Horticultural Science
Min, X.J. & Bartholomew, D.P., 1996. Effect of plant growth regulators on ethylene production, 1-aminocyclopropane-1-carboxylic acid
oxidase activity, and initiation of inflorescence development of pineapple. J. Plant Growth Regul. 15 (3), 121-128.
Parton, E., 2001. Flower biology and crossing barriers in Bromeliaceae. Doctoral thesis Catholic University of Leuven, Leuven.
Plever, H., 2004. Success with ethylene pills - the end of a long history using chemicals to induce bloom. In: Plever, H. (Ed.), Bromeliana 41
(12), 1-3.
Py, C., Lacoeuilhe, J.J., Teisson, C., 1984. L'ananas: sa culture, ses produits. G.P. Maisonneuve & Larose, Paris.
Rohrbach, K.G. & Taniguchi, G., 1984. Effect of temperature, moisture and stage of inflorescence development on infection of pineapple by
Penicillium funiculosum and Fusarium moniliforme var. subglutinans. Phytopathology 74 (8), 995-1000.
Turnbull, C.G.N., Sinclair, E.R., Anderson, K.L., Nissen, R.J., Shorter, A.J., Lanham, T.E., 1999. Routes of ethephon uptake in pineapple
(Ananas comosus) and reasons for failure of flower induction. J. Plant Growth Regul. 18, 145-152.
Yang, S.F., 1969. Ethylene evolution from 2-chloroethylphosphonic acid. Plant Physiol. 44, 1203-1204.
Table 1. The ethylene sources, with and without activated carbon, on the percentage of plants at the same flowering stage 73 days
after treatment.
Treatment
Ethylene, g
Activated carbon
Flowering (%)
Flowering stage
Control
0
6.7 ± 2.0 (A)*
No flowering
Zeothene
0
92.7 ± 3.6 (A)
3rd flower
Ethephon
0
82.6 ± 3.3 (AB)
2nd flower
Ethylene spray 0.389
2.86 g/L
78.4 ± 7.1 (B)
3rd flower
Ethylene
0.389
2.86 g/L
90.6 ± 4.6 (AB)
3rd flower
0.389
0
83.0 ± 11.7 (AB)
3rd flower
0.292
0
89.6 ± 5.2 (AB)
3rd flower
0.195
2.86 g/L
93.2 ± 7.2 (A)
3rd flower
0.195
0
95.0 ± 7.6 (A)
3rd flower
*Treatments followed by the same letter were not significantly different (*P < 0.05) from each other. (n = 100; repetitions = 3).
Figure 1. Ethylene gas absorption rates into pure water and water-activated
carbon solutions (n = 3). Significant differences (P<0.05) between the 5 %
solution and the other solutions are indicated by asterisks above the data points.
9
Newsletter of the Pineapple Working Group, International Society for Horticultural Science
MD-2 Pineapple Plant Maturity and Flowering
Van de Poel B.1 *, Ceusters J.2, De Proft M.P.2
1 BIOSYST-MeBioS, Department of Biosystems, Faculty of Bioscience Engineering, Katholieke Universiteit Leuven, W illem de
Croylaan 42, B-3001 Leuven, Belgium
2 Division of Crop Biotechnics, Department of Biosystems, Faculty of Bioscience Engineering, Katholieke Universiteit Leuven,
W illem De Croylaan 42, B-3001 Leuven, Belgium. E-mail of corresponding author: bram.vandepoel@biw.kuleuven.be
Flowering induction is a crucial aspect of pineapple cultivation. Growers all around the world treat their plants with
ethylene, or an ethylene releasing compound, to obtain simultaneous flowering. This practice will result in a homogeneous flower
and fruit development (Min & Bartholomew, 1996). It is important to apply the ethylene treatment at the right moment. Young
plants will produce smaller fruits, while older plants are more subjected to natural flowering, which disturbs flowering
homogeneity (Kerns, 1939). It is crucial to monitor plant development. This is done by sampling D leaves over a period of time.
D-leaf fresh weight or nutrient status are commonly used parameters to monitor plant development (Malézieux et al., 2003). The
optimal time for flowering induction, is the moment when plants have reached full biomass and have not yet started flowering due
to natural induction. Although biomass can be monitored accurately in individual fields, the actual maturity stage of the plants is
mostly unknown. In the literature this topic has been discussed but not yet fully determined. Das et al. (1965, cited by Norman,
1982) stated that pineapple plants should have at least 21 leaves before they are susceptible to flowering induction. Py et al.
(1984) suggested that plants ('Smooth Cayenne') should weigh at least 1 kg before applying flower induction treatments.
W e performed a field trial to find out at which exact stage during development, pineapple plants were completely
susceptible to an external ethylene treatment. Our experiment was conducted on a commercial pineapple plantation in Ecuador
(Quevedo) during the months of July and August 2006. The plantation is a monoculture of the new MD-2 hybrid. Plants of
different physiological age, starting from 1 month after planting (MAP) up to 8 MAP were given an ethylene treatment. The
different age groups were grown on different fields due to optimal management practices. All plants originated from shoots or
suckers (450 - 500 g) and were treated by standard cultural practices such as irrigation, fertilization and pest control to obtain
optimal plant growth and maximal fruit production. All plants used in the experiment were induced at the same moment to prevent
variation in weather conditions, which can influence the flowering induction treatment. The ethylene treatment was done by
dropping 3-4 zeothene pills in the central cup of each plant (n = 25). Zeothene is a new flowering induction agent (Van de Poel et
al., Pineapple News, this issue; Van de Poel et al., Scientia Horticulturae, 2009). Briefly, it is a zeolite pearl containing p ure
ethylene gas, which is released upon contact with the water standing in the central cup of the plant. One month after flowering
induction, the apical meristem was visually examined by making a longitudinal stem section. A vegetative apical meristem is
characterized by a flat dome shape, while a generative plant shows a vertical elongation of the apical meristem (Kerns et al.,
1936). Plant characteristics such as plant fresh weight (without roots) (g), number of leaves (including invisible young leaves),
D-leaf length (cm) and D-leaf fresh weight (g) were also recorded one month after flowering induction.
At one month after treatment, plants one MAP were not susceptible to external ethylene while at 2 MAP only 27 % of the
plants showed flowering (Table 1). At two MAP plants had a flowering stem 1.07 ± 0.18 cm long. At 3 MAP and older MD-2
plants were completely sensitive (100 %) to an external ethylene treatment. The average length of the flowering stem at one month
after treatment was not significantly different for plants of 3 MAP up to 8 MAP (3.10 - 3.53 cm). The growth of the flowering
stem was independent of plant size at one month after induction for plants at 3 MAP or later while 2 MAP plants were less
sensitive to ethylene forcing and had retarded flower development. These results suggest that plants of 3 MAP are fully mature.
Nevertheless, natural flowering is mostly observed much later during plant development. This might indicate that ethylene
perception is different for external applied ethylene and de novo synthesized ethylene due to natural flowering conditions.
Additional research on ethylene production and ethylene perception of the apical meristem, in relation to plant development,
should bring forth new insights in flowering sensitivity of pineapple.
During the first three months after planting, the young plant will mainly invest in rebuilding damaged tissue and root
development as they adapt to their new environment. During this period, plant growth, D-leaf growth and new leaf development
are limited (Table 1, Figure 1-4). The data are presented using box plots where the smallest observation is shown by a short bar,
usually below the box and easily visible in Figures 3 and 4. The lowest horizontal bar forming the base of the box illustrates the
lower quartile (25%), the second, the median (50%), the third, the higher quartile (75%), and the largest observation is indicated
by a short bar above the box, also easily visible in Figures 3 and 4. The mean is the filled circle in the middle of the box and the
median and the mean are equal when the middle bar in the box falls on the filled circle.
After the initial lag phase, growth in terms of plant fresh weight (Figure 1) and D-leaf development (length and fresh
weight) (Figure 3 & 4) becomes exponential. The total number of leaves increased linearly after the initial lag phase (Figure 2). At
around 8-9 MAP when the optimal plant weight is reached, vegetative development, including new leaf development, ceases due
to ethylene forcing. It is interesting that actual physiological maturity in terms of susceptibility to ethylene forcing coincides with
10
Newsletter of the Pineapple Working Group, International Society for Horticultural Science
the end of the initial lag phase (3 MAP). These results show that young pineapple plantlets first have to outgrow an initial phase
(recovery/adaptation) before they are susceptible towards an external ethylene treatment.
W e can conclude that MD-2 hybrid pineapple plants reach physiological maturity by 3 MAP even though they are not yet
large enough to produce a marketable fruit. W e can also conclude that D-leaf length and fresh weight are reliable parameters
representing general plant growth and can be used to determine the actual plant maturity border.
Table 1: Percentage of flowering and length of the flower stem one month after applying zeothane ethylene carrier and D-leaf
length, D-leaf fresh weight, plant fresh weight and leaf number for MD-2 pineapple plants (n = 25) at one to eight months after
planting.
Planting Flowering Flower stem
D-leaf
Plant
time
%
length (cm)
Length (cm)
Fresh weight (g)
Fresh weight (g)
Nr of leaves
1 MAP
0
59.7 ± 6.1 (A)
26.8 ± 5.8 (A)
354 ± 49.3 (A)
23.2 ±1.6 (A)
2 MAP
27
1.07 ± 0.18 (A)
57.0 ± 4.2 (A)
24.6 ± 4.6 (A)
443 ± 71.12 (A)
24.4 ± 3.0 (A)
3 MAP
100
3.10 ± 0.95 (B)
58.8 ± 5.1 (A)
24.6 ± 5.0 (A)
630 ± 90.2 (A)
29.2 ± 3.8 (A)
4 MAP
100
3.51 ± 0.32 (C)
73.8 ± 6.3 (B)
46.8 ± 8.5 (B)
829 ± 172.8 (A)
32.0 ± 4.1 (A)
5 MAP
100
3.53 ± 0.28 (C)
94.0 ± 7.5 (C)
70.4 ± 16.1 (C)
1338 ± 203.6 (B)
35.4 ± 4.5 (A)
6 MAP
100
3.33 ± 0.17 (BC) 105.2 ± 6.9 (D) 94.2 ± 15.9 (D)
1965 ± 406.8 (C)
44.2 ± 2.3 (B)
7 MAP
100
3.47 ± 0.21 (C)
111.3 ± 5.7 (E) 113.6 ± 19.2 (E)
2161 ± 328.8 (C)
42.0 ± 4.7 (B)
8 MAP
100
3.45 ± 0.16 (C)
120.5 ± 7.5 (F) 141.6 ± 17.9 (F)
2817 ± 253.7 (D)
48.0 ± 2.6 (B)
References
Kerns, K.R. 1936. Field results of acetylene on pineapple plants. Pine Quarterly 6, 95-114.
Kerns, K.R., Collins, J.L., Kim, H., 1939. Developmental studies of the pineapple Ananas comosus (L.) Merr. I. Origin and growth of
leaves and inflorescence. New Phytol. 35 (4), 305-317.
Min, X.J., Bartholomew, D.P., 1996. Effect of plant growth regulators on ethylene production, 1-aminocyclopropane-1-carboxylic acid
oxidase activity, and initation of inflorescence development of pineapple. J. Plant Growth Regul. 15 (3), 121-128.
Malézieux, E., Côte, F., Bartholomew, D.P., 2003. Crop environment, plant growth and physiology. In: Bartholomew, D.P., Paull, R.E.,
Rohrbach, K.G. (Eds.), The Pineapple Botany, Production and Uses. CAB International, Wallingford, pp. 69-107.
Norman, J.C., 1982. Growth, Flowering and Fruiting of 'sugerloaf' Pineapple, Ananas comosus (L.) Merr., as Influenced by Growth
Regulators and Cultural Practices. Rheinischen Friedrich-Wilhelms-Universität Bonn, Bonn.
Py, C., Lacoeuilhe, J.J., Teisson, C., 1984. L'ananas: sa culture ses produits. G.P. Maisonneuve & Larose, Paris.
Van de Poel, B., Ceusters, J., De Proft, M.P., 2009. Determination of pineapple (Ananas comosus, MD-2 hybrid cultivar) plant maturity,
the efficiency of flowering induction agents and the use of activated carbon. Sci. Hort. 20, 58-63.
Figure 1 (left). Box plots of the evolution of ‘MD-2' pineapple plant fresh weight from planting to time of flower induction at
8 months after planting. Figure 2 (right). Box plots of total leaf number of ‘MD-2' pineapple plants from planting (0 months
after planting, MAP) to 11 MAP.
11
Newsletter of the Pineapple Working Group, International Society for Horticultural Science
Figure 3 (left). Box plots of D-leaf length (cm ) of ‘MD-2’ pineapple plants from planting (0 m onths after
planting, MAP) to harvest at 12 MAP. Figure 4 (right). Box plots of D-leaf fresh weight of ‘MD-2’ pineapple plants from
planting (0 months after planting, MAP) to harvest at 12 MAP.
News From Brazil
New Records of Scale Insect Pests of Pineapple and Their Natural Enemies in the State of
Espírito Santo, Brazil
Mark P. Culik and José A. Ventura, Instituto Capixaba de Pesquisa, Assistência Técnica e Extensão Rural - INCAPER, Rua
Afonso Sarlo 160, CEP 29052-010, Vitória, Espírito Santo, Brasil. markculik3@yahoo.com
Scale insects (Hemiptera: Coccoidea) are important pests of pineapple and many other agricultural crops. Integrated pest
management (IPM) of such insects depends on knowledge of the pest species and natural enemies of the pests commonly present
in the crops of specific areas to obtain accurate information on the best management methods available. Unfortunately, relatively
little is known of the insect fauna, including scale insects and their natural enemies, in tropical areas such as the Brazilian State of
Espírito Santo (Culik et al. 2007, Culik et al. 2008). Therefore, surveys of scale insects and their natural enemies were conducted
in Espírito Santo to support development of integrated pest management as well as better document the biodiversity in this region.
Scale insects and their natural enemies were collected during surveys of the insect fauna of pineapple and when noticed on
plants during fieldwork or other activities in Espírito Santo in 2006 to 2008. Samples of plant parts (fruits, leaves, stems) infested
with scale insects were collected from locations ranging from municipalities of Serra in the north (20.13S; 40.31W ) to Anchieta in
the south (20.81S; 40.65W ) and Vitória (20.32S; 40.35W ) on the coast to municipalities in the interior of the state such as
Domingos Martins (20.38S; 41.050W ), from a variety of sites including experimental research plots, residences, and a
greenhouse. Samples were transported to the Espírito Santo rural research and extension institute INCAPER (Instituto Capixaba
de Pesquisa, Assistência Técnica e Extensão Rural) headquarters in Vitória for preservation and identification of the scale insects
and their natural enemies. Samples were also placed in plastic containers covered with cloth to allow development of natural
enemies present and examined every few days for several weeks to collect adult parasites and predators that emerged. Specimens
collected were sent to taxonomic specialists to confirm identifications when necessary.
Based on this research, four scale insect species that are potential pests of pineapple, Melanaspis smilacis, Unaspis citri,
Dysmicoccus brevipes, and Planococcus minor, were recorded for the first time in different municipalities in the State (Table 1).
In addition, a wide variety of natural enemies of the scale insect pests of pineapple were collected, including 5 new species of
predators that are currently being described, and 6 species of parasitoids that are also being studied further to confirm
identifications (Table 2).
Most of the scale insect species collected in this study are polyphagous and widely distributed (Ben-Dov et al. 2006). Thus,
they are potential pests of many agricultural crops in many tropical areas. However, natural enemies of many of these scale insects
were commonly found associated with these scale insects in Espírito Santo indicating the importance of using IPM methods, and
avoiding improper and harmful management practices such as misuse of pesticides, to prevent destruction of beneficial insects and
natural enemies of scale insects that may commonly help control scale insect and other pests in this area.
12
Newsletter of the Pineapple Working Group, International Society for Horticultural Science
Acknowledgements
Research Support provided by the Fundação de Apoio à Ciência e Tecnologia do Espírito Santo - FAPES, FINEP, and CNPq.
Literature Cited
Ben-Dov, Y., D. R. Miller, and G. A. P Gibson. 2006. ScaleNet. Available at: http://www.sel.barc.usda.gov/scalenet/scalenet.htm. 25 Jul 2008.
Culik, M. P., D. S. Martins, J. A. Ventura, and V. F. Wolff. 2008. Diaspididae (Hemiptera: Coccoidea) of Espírito Santo, Brazil. J. Insect Sci.
8(17):1-6.
Culik, M. P., D. S. Martins, J. A. Ventura, A. B. G. Peronti, P. J. Gullan, and T. Kondo. 2007. Coccidae, Pseudococcidae, Ortheziidae, and
Monophlebidae (Hemiptera: Coccoidea) of Espírito Santo, Brazil. Biota Neotropica 7:61-65.
Table 1. New records of scale insect
Taxon
DIASPIDIDAE
Melanaspis smilacis (Comstock)*
Unaspis citri (Comstock)
PSEUDOCOCCIDAE
Dysmicoccus brevipes (Cockerell)*
pests (Hemiptera: Coccoidea) of pineapple in the State of Espírito Santo, Brazil (2006-2008).
Municipality
Associated crop/plant
Note
Planococcus minor (Maskell)*
Domingos Martins
Domingos Martins
Ananas comosus
Citrus sp.
New register in municipality
New register in municipality
Serra
Domingos Martins
Vitória
Cocos nucifera
Ananas comosus
Syzygium jambos
New host record in State
New register in municipality
New host record in State, new register
in municipality
*Associated natural enemies (parasitoids and predators) also collected.
Table 2. New records of predators (Diptera: Cecidomyiidae, Drosophilidae) and parasitoids (Hymenoptera: Chacidoidea) of scale
insect pests of pineapple in the State of Espírito Santo, Brazil (2006-2008).
Taxon
Municipality
Associated crop/plant
Associated scale insects
CECIDOMYIIDAE
Diadiplosis sp. nov.1
Vitória
Syzygium jambos (fruits)
Planococcus minor, Planococcus halli
Diadiplosis sp. nov.2
Sooretama, Cachoeiro
Ananas comosus (seedlings, Dysmicoccus brevipes
do Itapemirim, Domingos
fruit, leaves and crowns)
Martins
Diadiplosis sp. nov.3
Domingos Martins
Ananas comosus (fruit)
Pseudococcus cf. jackbeardsleyi;
Coffea arabica
(mummies), Saissetia cf. coffeae
(fruits, stems)
Diadiplosis sp. nov.4
Domingos Martins
Coffea arabica
Pl. cf. citri,
(fruits, stems)
Ps. cf. jackbeardsleyi,
S. cf. coffeae
DROSOPHILIDAE
Rhinoleucophenga sp. nov.
Cachoeiro do Itapemirim
Ananas comosus (fruit)
Dysmicoccus brevipes
APHELINIDAE
Encarsia cf. aurantii
Domingos Martins
Ananas comosus
Diaspis boisduvalii
Encarsia lounsburyi
Domingos Martins
Ananas comosus
Melanaspis smilacis
ENCYRTIDAE
Adelencyrtus modestus
Sooretama
Ananas comosus
Diaspis boisduvalii and Melanaspis
smilacis
Sooretama;
Ananas comosus:
Dysmicoccus brevipes
Anagyrus cf. cercides
Serra
Cocos nucifera
cf. Anagyrus
Cachoeiro do Itapemirim;
Ananas comosus;
Dysmicoccus brevipes;
Jaguaré
Coffea canephora
cf. Ferrisia and Pseudococcus
cf. Hambletonia
Sooretama;
Jaguaré
Ananas comosus;
Coffea canephora
Dysmicoccus brevipes;
cf. Ferrisia and Pseudococcus
cf Leptomastix
Jaguaré;
Domingos Martins
Sooretama
Coffea canephora;
Coffea arabica
Ananas comosus
cf. Ferrisia and Pseudococcus;
cf. Planococcus
Dysmicoccus brevipes
Serra
Cocos nucifera
Dysmicoccus brevipes
Prochiloneurus sp.
EULOPHIDAE
Diglyphomorpha sp.
—
13
Newsletter of the Pineapple Working Group, International Society for Horticultural Science
News From Cuba
Cryogenic Strategy For The Establishment Of Pineapple (Ananas Comosus L. Merrill)
Germplasm Bank At Bioplantas Centre (Cuba)
Martinez-Montero M.E.*, Mendez-Pelegrin R. & Martinez J.
Bioplantas Centre, Carretera a Morón Km 9 CP 69450.University of Ciego de Ávila, Ciego de Ávila. Cuba. Teléf:(5333)224016/225768. Fax: (53-33)266340. *e-mail for correspondence: marcosem@bioplantas.cu
Cryopreservation of pineapple tissue has been based on protocols using the vitrification procedure. However, further research
is necessary to identify the different technical factors required to obtain the appropriate cryogenic strategy for routine application
to a wide number of genotypes. Moreover, the visualization of structural changes during the development of a cryopreservation
procedure for pineapple has not been accomplished until now. For the above reasons in the present research different key technical
issues were determined during the establishment of a cryogenic strategy to induce dehydration tolerance to a highly concentrated
vitrification solution to improve the survival rates for in vitro grown shoot tips of pineapple after immersion in liquid nitrogen
(LN). The best established conditions were: type of shoot tip (consisted in meristematic dome area and 3-4 primordial leaves 2.5 –
3 mm in size); 2 days of preculture in 0.3 mol L-1 sucrose; application of the loading solution (0.4 mol L -1 sucrose + 2 mol L -1
glycerol) for 25 min at 25°C; 7 hours of dehydration at 0°C with plant vitrification solution number three (PVS3: 50% w/v
glycerol + 50% w/v sucrose). W ith these best conditions, histological analysis of the structural changes of cryopreserved pineapple
shoot tips revealed that only cells localized in the meristematic area and in young leaf primordia had a few cellular alterations
while their morpho-physiological characteristics remained almost intact. Moreover, the vitrification procedure was successfully
applied to nine accessions of the in vitro collection at Bioplantas Centre. These results constitute a very important step in the
validation of cryopreservation protocols for pineapple germplasm conservation and the real establishment of its cryobank.
Phenotypic Characterization of Field-grown Pineapple Transgenic Plants
Lourdes Yabor* , Bárbara Valle, Carol Carvajal, Carlos Aragón, Martha Hernández, Justo González, Marcos Daquinta, Ariel
Arencibia & José Carlos Lorenzo
Laboratory for Plant Breeding, Bioplant Center, University of Ciego de Avila, 69450, Cuba (*Fax: 53 33 266340, E-mail:
lyabor@bioplantas.cu)
W e previously introduced the bar gene, along with chitinase and AP24 genes, into the pineapple genome. The present report
focuses on the phenotypic evaluation of the first vegetative generation of transgenic plants. Three plant materials were compared:
macropropagated controls (non-transformed), micropropagated controls (non-transformed), and micropropagated transformed
plants. From each group, 50% of the plants were sprayed with FINALE ® 3 months after initiation of the experiment. The
phenotypic characterization was performed after one year of field growth. FINALE ® killed all non-transgenic plants.
Micropropagated transformed plants sprayed with FINALE® , did not show phenotype differences from micropropagated
transformed plants not sprayed with the herbicide. Between the micropropagated transformed plants sprayed with FINALE ® and
the micropropagated control plants not sprayed, there were two experimental differences: the genetic transformation and the
herbicide application. The combined effects of these two factors caused modifications in levels of phenolics (cell wall-linked,
free, total) and proteins. Moreover, they changed the fruit mass without crown. Between the micropropagated transformed plants
sprayed with FINALE® and the macropropagated control plants not sprayed, there were three experimental differences: the genetic
transformation, the herbicide application, and the in vitro culture. They provoked changes in levels of chlorophylls (b, total) and
proteins. Furthermore, activities of phenylalanine ammonia – lyase, superoxide dismutase and glutamine synthetase -were
modified. The plant height and diameter, and the crown height were also changed by these three experimental differences. Until
now we have evaluated transformed pineapple plants during hardening and field growth. Although some unexpected variations
were recorded, we believe that they are not relevant enough to justify rejection of transgenesis as an important tool for pineapple
genetic improvement.
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Newsletter of the Pineapple Working Group, International Society for Horticultural Science
Proteases Expressed in Response to in Vitro Culture of Pineapple (Ananas comosus (L.)
Merr.)
Hernández M 1 , Pérez A1 , Lorenzo J2, Carvajal C 1, Mora M 1, Natalucci C 3, Jorrín J. 4
Laboratorio de Ingeniería Metabólica. Centro de Bioplantas. Universidad de Ciego de Ávila, Ciego de Ávila, CP 69450, Cuba.
E-mail: mhernandez@bioplantas.cu.
2
Laboratorio de Mejoramiento Genético. Centro de Bioplantas. Universidad de Ciego de Ávila, Ciego de Ávila, CP 69450, Cuba.
3
LIPROVE, Departamento de Ciencias Biológicas, Facultad de Ciencias Exactas, Universidad Nacional de la Plata. La Plata.
Argentina.
4
Departamento de Bioquímica y Biología Molecular. Universidad de Córdoba. Córdoba. Campus de Rabanales, Edificio Severo
Ochoa (C6), 14071 Córdoba, España.
1
Biotechnology has become an important tool to produce proteases. Bromeliaceae family plants usually contain high
concentration of thiol proteases. Although pineapple plants have been found to produce proteases, most of the biotechnological
investigations on this crop have been focused on propagation. Plant tissue culture techniques have provided many solutions to
basics questions and practical problems in plant biology. Therefore, considerable attention has been focused on the possibility of
applying efficient plant tissue culture methods to physiologically active enzymes isolation. W e decided to modify pre-elongation
phase during pineapple micropropagation in temporary immersion bioreactors (TIB), looking for an increase of protease
excretion. Seven experiments were performed to evaluate the effects of culture duration, levels of gibberellic acid (GA), 6benzyladenine (BA), different levels of sucrose, inorganic salts, inositol and thiamine. The following indicators were recorded:
shoot fresh mass per bioreactor; and protein concentration, proteolytic activity, and specific protease activity in culture media.
Specific protease activity was highest at 21 d of culture, with 90g/L of sucrose, 4.2 µmol/L of GA, 100% MS salt strength, 0.1
mg/L of thiamine. Moreover, BA and inositol produced a negative effect. Proteases expression in response to in vitro culture by
2D- electrophoresis was evaluated. W e found proteolytic activity in pineapple shoots cultured in vitro. The highest specific
protease activity was recorded in shoots cultured in TIB. Multiplication phase in vitro did not cause a remarkable protease
production in shoots. Proteome of shoots cultured in different in vitro phase were compared. Molecular mass of some protein
spots were between 21 500- 31 000 Da. This parameter was similar to those indicated for cysteine proteases from Bromeliaceae.
A protease was detected in TIB culture media. Retention time in RP-HPLC and molecular mass of the major protein detected in
TIB culture media showed high similarity to stem bromelain.—
News From France
The Domestication of Pineapple: Context and Hypotheses
Geo Coppens d’Eeckenbrugge, CIRAD, UMR 5175 CEFE, 1919 Route de Mende, 34293Montpellier, France
geo.coppens@cirad.fr
Marie-France Duval, CIRAD, UPR Multiplication Végétative, Avenue Agropolis, TA A-75/03, 34398 Montpellier,
Francemarie-france.duval@cirad.fr
Introduction
The present economic importance of pineapple is easily justified by its unique characteristics as a fruit, which ensured its very
rapid diffusion and adoption, first at the continental level in all of tropical and subtropical America, and later at the global level in
all tropical areas. However, as with other major crops of Amazonian or peri-Amazonian origin (e.g., cassava), how these unique
characteristics were developed in pre-Columbian times, i.e., the process of its evolution under cultivation or domestication, has
been poorly investigated. The reason for this must be mostly sought in the relatively limited interest that has long prevailed for the
domestication of vegetatively propagated crops, particularly fruit crops. There also has been limited interest or limited study of the
prehistory of the humid tropics, because there is poor conservation of archaeological macro-remains in the region. On the other
hand, the views of archaeologists on crop domestication in relation to the birth of agriculture and the evolution of human societies
have evolved considerably in the last decade. The greater acceptance within the scientific community of an Amazonian cradle of
plant domestication and agriculture provides a favourable context for a new evaluation of the pineapple case. W ithin that context,
we review here the results of recent research on pineapple genetic resources and the available archaeological data in an attempt to
trace the path of development of this impressive fruit.
Crop domestication and its archaeological context
Domestication is a form of co-evolution in which humans and their crops and animals become dependent upon each other.
The domestication syndrome is a set of morphological, phenological and physiological traits that are modified by conscious or
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Newsletter of the Pineapple Working Group, International Society for Horticultural Science
unconscious human selection, and cause this dependence. A classical example is that of cereals, whose cultivation was first based
on seeds from the wild. The fixation of the domesticated forms was slow even in these annual crops, requiring more than one
millennium before they constituted most of the cultivated populations (Tanno and W ilcox, 2006; Fuller, 2007; Fuller et al., 2009).
The process of crop domestication and the identification of centres of crop origin are of particular interest to plant breeders
because of the importance of wild relatives and landraces as genetic resources for the improvement of modern crops and their
adaptation to marginal growing conditions. W ith regard to domestication in the humid tropics, there was the perception of the
tropical forest as a hostile habitat for the development of human societies and agriculture. W ith few exceptions (e.g., Carl Sauer),
scientists thought that forest dwellers were relative latecomers to agriculture, first practicing agriculture between 4000 to 5000
years before the present (BP). W hile the explanatory model for the so-called Neolithic Revolution involving the development of
cereal crops was extended to temperate and subtropical Asia, the evolution of civilization and agriculture in the two main rain
forest areas, the Amazon and the Congo basins, long remained at best poorly studied. The idea that these reservoirs of
megadiversity remained unexploited for agricultural development was long held by followers of Vavilov and Harlan (e.g.,
Hawkes, 1998), prominent among the early scientists proposing theories of crop domestication. Elucidation of crop domestication
in the humid tropics was also delayed by the strong focus on major species propagated by seeds, mostly cereals and pulses, and the
relative neglect of vegetatively propagated and perennial crop plants. Even in the Fertile Crescent, relatively little attention has
been paid to fruits like figs, despite the antiquity and importance of their cultivation and their potential as botanic/genetic markers
of human movements in the Near East and the Mediterranean. In fact, horticulture/vegeculture may even have preceded cereal
cultivation (Kislev et al., 2006).
The thinking on plant domestication and the advent of agriculture has changed considerably in recent years, with new
scenarios involving the humid tropics and non-cereal plants. There is evidence of ancient major crop cultivation in the tropical
lowlands, particularly New Guinea and/or Melanesia (root crops, bananas and sugar cane; see Neumann, 2003) and the
Neotropics. The Amazon and its periphery have been recognized as the cradle of ancient complex civilizations (Gibbons, 1990)
and domestication of cassava (Olsen and Schaal, 2001), two species of chilli pepper (Capsicum baccatum and C. chinense), jack
bean (Canavalia plagiosperma), arrowroot (Maranta arundinacea), cocoyam (Xanthosoma sagittifolia), llere´n (Calathea
allouia) (Piperno and Pearsall, 1998, cited in Olsen and Schaal, 2001 and Pickersgill 2007), pineapple (Leal and Coppens
d'Eeckenbrugge, 1996), cocoa (Motamayor et al. 2002), and peach palm (Bactris gasipaes; Clement, 1995). Archaeological
evidence, including 7000 years old pottery (Roosevelt et al., 1991), vast extensions of fertile anthropogenic soils (Amazonian dark
earths; W oods et al. 2009) and topographical modifications (Heckenberger et al., 2003), indicates the very early development of
agriculture in this region, leading to the establishment of significant populations (estimated between 2 and 5 million at the time of
European contact; Denevan, 1992; Hornborg, 2005), and significant forest transformation through enrichment or degradation by
slash and burn agriculture (Piperno, 2006; but see also Bush et al. 2007). The Amerindian occupants of Amazonia made their
living by hunting, fishing and the cultivation or management of more than 138 plant species (Clement, 1999), among them a very
large number of fruit trees (Miller and Nair, 2006). Combining mobile and sedentary strategies (see Rival, 2006), they managed
the forest in such a way as to encourage the concentration of useful species, in processes accompanied by selection of superior
genotypes of fruit tree species (Gnecco, 2003). Crop domestication appears to be quite as old as these concentration/improvement
processes, and long distance exchanges ensured an early diffusion of many crops between Amazonia, the Andes and the Pacific
Coast, as well as between South America and Mesoamerica (Stone, 1984). Indeed, cassava was present in northern Colombia by
7500 BP, in Panama by 7000 BP (Zeder et al., 2006), which compares well with the presence of maize, of Mesoamerican origin,
in South America by 7500-7800 BP, according to data from Colombia and southwest Ecuador (Pohl et al., 2007; Dickau et al.,
2007). Thus, cassava was probably domesticated roughly at the same time as maize, i.e. in an 8-10,000 BP interval (9000 BP
according to genetic data of Matsuoka et al., 2002), after a long phase of wild type cultivation. This timeframe is also consistent
with data for squash, which seemed to precede maize, being domesticated around 10,000 BP in Mexico (Cucurbita pepo; Smith,
2001), Ecuador and Peru (C. moschata ; Piperno and Stothert, 2003 ; Dillehay et al., 2007).
Pineapple domesticates and their wild relatives
In seed-propagated crop species, the reproductive modifications induced by domestication often result in partial or complete
reproductive isolation, i.e. the domesticate becomes a new species. These modifications can be so great that the identification of
the crop wild relatives is problematic. Even without a sexual barrier, maize looked so distinct from its teosinte genitor, which may
grow as a weed in the same fields, that the two forms of Zea mays were once classified in different genera.
No such difficulty appears in the case of the pineapple, and domestication has not produced any clear, qualitative,
morphological or physiological differentiation, or reproductive isolation. In fact, the genus Ananas only includes two species, the
pineapple, A. comosus (L.) Merril, and the gravata or yvira, A. macrodontes Morren (Coppens d'Eeckenbrugge and Leal, 2003).
The former is a normally diploid species (2n = 50) that includes five botanical varieties, three of which are domesticates. Its
natural distribution includes all tropical South America east of the Andes. The latter is a self-fertile tetraploid (2n = 4x = 100),
whose inflorescence lacks a crown and vegetative reproduction is ensured by stolons. It grows wild in forests of southern South
America. Although exploited by natives for the production of fibres (Corrêa, 1952), it shows no sign of domestication. The two
species exhibit limited differentiation in molecular genetic studies (Duval et al., 2001 and 2003), however their ploidy difference
constitutes a clear reproductive barrier.
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Newsletter of the Pineapple Working Group, International Society for Horticultural Science
Cultivated pineapples
The three cultivated Ananas botanical varieties are A. comosus var.
comosus, the pantropical pineapple cultivated for its spectacular and
exquisite large fruit, A. comosus var. erectifolius, a small-fruited pineapple
cultivated for its fibre, and A. comosus var. bracteatus, a robust pineapple
with multiples uses, involving its medium-sized fruit for juice and its
armed leaves for fences. The two latter varieties are now increasingly
cultivated as ornamentals.
In A. comosus var. comosus, the syncarp grows very significantly after
anthesis, so the fruit is generally very large and fleshy (up to several
kilograms in certain cultivars; Figure 1), with many fruitlets ("eyes"); they
are borne on a wide and strong, relatively short, peduncle. Seeds are rare in
the fruits, because of reduced fertility, conjugated with stronger
self-incompatibility and monoclonal cultivation (Coppens d'Eeckenbrugge
et al., 2003). Vegetative reproduction, through shoots, in the vernacular of
pineapple, slips, suckers and crowns, is often initiated after floral
induction; under tropical conditions, these propagules tend to resume
growth after fruit maturity. The plant has numerous wide leaves (40-80),
with antrorse spines; these marginal spines are generally smaller and
denser than in wild varieties and can be partially or completely suppressed
by dominant mutations.
At the time of the Conquest, A. comosus var. comosus was planted
throughout tropical America, and included cultivars having wide variation
in fruit size, shape, colour, and flavours. Considerable morphological and
genetic diversity was found in the western Amazon and the eastern
Guianas (Duval et al., 1997, 2000, 2003). Variation in adaptation to
different environments, including Andean hillsides was also evident. The
fruit was widely consumed, and appreciated in the form of fermented
Figure 1. A 10 year-old young native proudly showing
a large pineapple from an Amazonian cultivar
drinks. Other traditional uses were based on its properties as a digestive,
(Photograph G. Coppens).
vermifuge, antiamoebic, abortifacient and emmenagogue, most of which
are related to the presence of a proteolytic enzyme complex in pineapples
as well as in many other bromeliads (Leal and Coppens d'Eeckenbrugge, 1996; Patiño, 2002).
Plants of A. comosus var. erectifolius are much less massive, with abundant and early shoots, frequent crownlets at the base of
the main crown, numerous erect, fibrous leaves and a small, very fibrous, inedible fruit borne on a long and slender peduncle
(Figure 2). In some clones, the fruit appears to be rare. A. comosus var. erectifolius is quite similar to the wild A. comosus var.
ananassoides except for its smooth leaves, a trait which is under monogenic control (Collins, 1960). A. comosus var. erectifolius is
not known to occur in the wild. It was cultivated in the W est Indies at the time of Conquest, and it is still cultivated by the natives
in the Guianas, including the Orinoco basin, and in the north of the Amazon basin, for the strong and long fibres associated with
its typical erect habit. Indeed, the dry fibres constitute 6% of the plant weight. They are used to make hammocks and fishing nets
(Leal and Amaya 1991), but now suffer competition from synthetic fibres and nylon. Vernacular names include curagua, curauá,
curaná, kulaiwat, and pitte. The typical absence of spines along the leaf margin, as well as its erect habit, is the likely result of
artificial selection for high yield of easily extractable fibres among strains of A. comosus var. ananassoides. Variety erectifolius
has recently found a new economic use in the production of cut flowers.
Ananas comosus var. bracteatus is an assemblage of two cultivated forms that show the same geographic distribution as A.
macrodontes, and that are morphologically and genetically intermediate between A. comosus and A. macrodontes (Figure 3). The
most common one corresponds to A. bracteatus sensu Smith & Downs, which was cultivated as a living hedge and harvested for
fibre and fruit juice, or for traditional medicine, in southern Brazil and Paraguay (Bertoni 1919). Indeed, its dense, long and wide
leaves are strongly armed by large antrorse spines, forming impenetrable barriers. It is very robust and still thrives in aband oned
plantations, but it seems unable to colonize new habitats. The syncarp is of intermediate size (0.5 to 1.0 kg), borne by a strong
peduncle, and covered by long and imbricate floral bracts, as in A. macrodontes. These bracts are bright pink to red at anthesis, to
within-cultivar variations (Duval et al. 2001, 2003) and suggesting a very narrow origin, possibly a single genotype. The second
producing a spectacular inflorescence. Morphological and genetic variations are very limited in this first form, being comparable
form, corresponding to A. fritzmuelleri Camargo, shares an additional trait with A. macrodontes, as it exhibits retrorse spines on
17
Newsletter of the Pineapple Working Group, International Society for Horticultural Science
Figure 2A. A. comosus var. erectifolius under cultivation for fibre in the
Amazon (Rio Negro basin; photograph of M.F. Duval), and Figure 2B,
as an ornamental, for export, in Côte d'Ivoire (photograph of G.
Coppens).
Figure 3. A. comosus var. bracteatus, a remnant from an old fence in a southern Brazil farm
(machete handle provides scale; photograph of M.F. Duval), and a variegated mutant used as a
garden ornamental in Martinique, FWI (photograph of G. Coppens).
18
Newsletter of the Pineapple Working Group, International Society for Horticultural Science
the leaf base. According to Camargo (1943) and Smith and Downs (1979), it was also used in living fences. It is a very rare form,
whose diversity has not been documented, only one clone being conserved in Brazil, by EMBRAPA and the botanical garden of
Rio de Janeiro. The chromosome number is 2n=2x=50 (Camargo, 1943).
Figure 4. Distribution of A. macrodontes (S) and A. comosus varieties ananassoides (A), parguazensis (P),
erectifolius (L), and bracteatus (B). Red roughly outlines the region of greatest morphological and genetic
diversity within A. comosus (including var. comosus and types intermediate between var. comosus and
ananassoides). Partial spininess is relatively frequent in this area. Green outlines an area with great
diversity of large-fruited clones (typical of var. comosus), where the "piping" leaf trait is relatively frequent.
Wild relatives of cultivated pineapples
W ild pineapple relatives include the varieties ananassoides and parguazensis of A. comosus and A. macrodontes.
W ithin A. comosus, wild botanical varieties display the highest genetic diversity, which is a common situation in crop gene
pools. The most common and diverse wild variety, A. comosus var. ananassoides, is also the likely ancestor of the cultivated
botanical varieties. It is generally found in savannahs or clear open forests, growing on soils with limited water-holding capacity
(sand dunes or "campinas", rocks, common on and around the Guiana shield) and forming populations of variable densities. In the
Guianas, it can also be found, although rarely, thriving in dense rain forest. In contrast, it is absent from the seasonally flooded
lands along the Amazon and its main southern tributaries, which seem to act as a barrier dividing its distribution in two main areas;
a northern one corresponding to the Guiana shield, Orinoco basin, and northern drainage of Rio Negro (i.e., from the Brazilian
19
Newsletter of the Pineapple Working Group, International Society for Horticultural Science
state of Amapá to eastern Colombia), and a southern one roughly corresponding to the Brazilian shield and north-eastern Brazil
(from the Brazilian states of Acre, Mato Grosso over to Pernambuco and down to Paraguay and northern Argentina) (Figure 4).
A. comosus var. ananassoides has long and narrow leaves, up to 2 m long and less than 4 cm wide, subdensely serrate with
wholly antrorse spines. The fruit peduncle is elongate (most often more than 40 cm) and slender (usually less than 15 mm wide).
In the southern part of its distribution, the inflorescence is very generally small, globose to cylindrical, and it shows little growth
after anthesis, so it has little flesh. The pulp is white or cream, very firm and fibrous, with high sugar content and acidity, and
numerous seeds. And its habitat appears mostly restricted to areas providing an open and markedly dry habitat (grass savannahs
and low open forests; Figure 5). In contrast, in the northern area, the habitats of A. comosus var. ananassoides appear more
variable (Leal and Medina, 1995), and a higher morphological diversity is observed, with clones producing larger, fleshy fruits (up
to 12-15 cm long) as the syncarp shows significant growth after anthesis. Their fruits were consumed in the Orinoco (Patiño,
2002) and are still occasionally consumed in the Guianas. Similar types, morphologically intermediate between the wild and
cultivated forms, are sometimes found in patches in secondary forest and savannahs in French Guiana, indicating an ancient
settlement, or cultivated in gardens (Figure 6). They constitute the most plausible basis for initial domestication in the Guianas.
Indeed, in the study of Duval et al. (2003), these intermediate phenotypes display four haplotypes, sharing three of them with A.
comosus var. comosus and all four with A. comosus var. ananassoides, which is consistent with the hypotheses of
semi-domestication or introgression between the two botanical varieties.
Figure 5. Typical habitat of A. comosus var. ananassoides, South of the Amazon basin in southern Brazil
(photograph of M.F. Duval) and inset, two clones from northern Mato Grosso (photographs G. Coppens).
The contribution of A. comosus var. parguazensis to the evolution of the cultivated pineapple is less likely, on geographic,
morphological, and genetic grounds. Its geographical distribution appears more centred, mostly corresponding to the basins of the
Orinoco and upper Rio Negro, the area of its greatest diversity, with a few observations in eastern Colombia and in north-eastern
Amazon (Coppens d'Eeckenbrugge et al. 1997; Duval et al. 2001, 2003). It grows in lowland forests, under canopies of variable
densities, from clearings or riverbanks to dense forest. As compared to specimens of A. comosus var. ananassoides growing in
close proximity, it seems restricted to shadier environments, because of lower water use efficiency (Leal and Medina 1995).
Morphologically, it differs from variety ananassoides by having wider leaves, slightly constricted at their base, and larger spines,
some of them retrorse (Figure 7). A few Orinoco/Rio Negro phenotypes appear to be intermediate between varieties parguazensis
and ananassoides, indicating some natural hybridization. However, retrorse spines and the basal leaf constriction have not been
observed in the cultivated pineapples. Instead A. comosus var. parguazensis appears genetically distinct, forming a particular
branch in the nuclear DNA as well as in the chloroplast DNA phylogenetic trees, with relatively few exceptions for the latter.
Duval et al. (2003) explain these exceptions as the result of hybridization with var. ananassoides in the Orinoco/Rio Negro region
and by a different genetic background for the rare specimens of eastern Guiana (which implies morphological convergence
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Newsletter of the Pineapple Working Group, International Society for Horticultural Science
between wild types of distinct origins). Four of the seven parguazensis chloroplast genotypes, including the most common ones,
are not shared with other botanical varieties (Duval et al. 2003). In conclusion, a contribution of A. comosus var. parguazensis to
the genomes of the cultivated pineapples cannot be ruled out, but it would be marginal, and necessarily indirect, through
occasional hybridization with A. comosus var. ananassoides as the wild ancestor of A. comosus var. comosus and A. comosus var.
erectifolius.
Figure 6. Habitats and morphology of A. comosus var. ananassoides, in French Guiana and Amapá state, Brazilian : A, B,
inselberg and rock savannah; E, intermediate phenotype with medium fruit on long peduncle in cultivation in wild lowland forest; C,
D, extreme range of fruits sizes commonly found (see pen at white arrows for scale) (photographs of G. Coppens).
The tetraploid A. macrodontes (Figure 8) only shows clear genetic affinity with A. comosus var. bracteatus, as they share rare
isozymes (García 1988) as well as nuclear DNA markers, and chloroplast DNA markers in the case of the former A. fritzmuelleri
Camargo (Duval et al., 2001, 2003). They also share most of their original geographic distribution in southern South America, and
several morphological traits such as wide leaves, strong spines, the presence of retrorse spines, fruit peduncles of intermediate
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Newsletter of the Pineapple Working Group, International Society for Horticultural Science
length and width, and bearing medium-size fruits with floral bracts longer than the individual flowers. On the other hand, A.
macrodontes is differentiated by the lack of a crown at the top of the syncarpic fruit and by vegetative reproduction by stolons.
The fruit flesh is low in acid and it contains numerous seeds. A. macrodontes appears to be highly self-fertile. The natural habitat
of A. macrodontes corresponds to humid forest areas, under semi-dense shade, from south-eastern Paraguay and north-eastern
Argentina up to Mato Grosso and coastal Brazil (Coppens d'Eeckenbrugge et al. 1997).
Figure 7 (left). A. comosus var. parguazensis (Rio Negro basin. Figure 8 (right). A. macrodontes in the wild
in southern Brazil (photographs of M.F. Duval).
Domestication syndrome in the cultivated varieties of Ananas com osus
The clearest modifications due to domestication in A. comosus var. comosus (Leal and Coppens d'Eeckenbrugge, 1996) are:
! Reduced susceptibility to natural flowering induction.
! A larger number of wider, and generally shorter, leaves.
! A wider and longer stem allowing a largergreater starch storage capacity.
! A significant increase in the number of flowers, correlated with a modification in their phyllotaxy.
! The enlargement of individual fruits (pineapple "eyes").
! A reduction in fruit fibrousness.
! Reduced seed production through the combination of lower sexual fertility and stronger self-incompatibility.
In the cultivars where the reduction of female fertility, i.e. the proportion of ovules producing a seed, is not very severe, it can
be counterbalanced by the higher number of flowers. In any case, as vegetative reproduction is largely dominant in Ananas, this
reduced sexual potential affects plant survival less than the changes in the vegetative organs and the plant vegetative cycle. Strictly
speaking, the domestication syndrome in A. comosus var. comosus lies in its lack of adaptation to the natural conditions prevailing
where the wild varieties are found. Pineapple plants from most cultivars can survive when their cultivation is abandoned, resisting
competition in sufficiently open vegetation and even in dry edaphic or climatic conditions. However, they do not propagate
efficiently to form subspontaneous feral populations. This may be due to the cost of an excessive harvest index (i.e., the
production of a relatively large fruit), limiting the capacity for vegetative propagation, and/or the loss of dispersal capacity, as only
man can transport large fruits and their crown over medium to long distances (assuming that no animal has an interest/capacity for
the dispersal of other vegetative propagules). Indeed, wild pineapple populations are distributed discontinuously in the Guianese
forests. They are most often found on relatively elevated areas (inselbergs, "rocky savannahs") where there is no risk of water
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Newsletter of the Pineapple Working Group, International Society for Horticultural Science
stagnation. Such sites are often isolated in the forest, which implies long distance vectors for seed dispersal, very probably large
birds and/or monkeys. Indeed, sexual propagation might play an important role in the initial foundation of wild populations, as
only one or two clones were observed at a given site (Coppens d'Eeckenbrugge, unpublished), while clones appeared distinct
among sites.
The situation appears to be similar for the cultivated botanical varieties A. comosus var bracteatus and A. comosus var
erectifolius, which do not show any capacity for spontaneous colonization in the wild. As in var. comosus, a large fruit size could
be limiting in var. bracteatus. For the small-fruited var. erectifolius, the loss of leaf spines probably increases its susceptibility to
herbivory, although leaf fibrousness might compensate. In addition, for those clones of var. erectifolius that rarely produce a fruit,
there is an additional restriction on sexual recombination, and thence for the plant adaptive potential, strengthening the
dependence on man. On the other hand, we must keep in mind that the main qualitative characteristic that distinguishes var.
erectifolius from var. ananassoides is the presence or absence of spines. Thus, when smooth-leaved clones of var. erectifolius
mutate back to the spiny condition, as has been observed in germplasm collections as well as under cultivation, these reverse
mutants should be formally classified in var. ananassoides. In this case, the domesticate status and the domestication syndrome
may look as fragile as the morphological difference with its wild ancestor.
How, where and when pineapple domestication may have proceeded?
Bertoni (1919) proposed that the pineapple was domesticated in southern South America by the Tupi-Guarani Indians who
would have diffused the crop in their northward migrations. Later, most reviewers of pineapple domestication (Collins, 1960;
Purseglove, 1972; Pickersgill, 1976; Sauer, 1993 ) accepted a southern origin. Only Brücher (1971), whose paper was written in
German and subsequently ignored, underlined the presence of wild forms and primitive cultivars in the north of South America
and proposed a Guianese origin. In any case, both hypotheses were based on very limited knowledge of pineapple diversity and
distribution. Leal and Antoni (1981) called attention to the greater morphological diversity to be found north of the Amazon.
Extensive expeditions in Venezuela, Brazil, and French Guiana (Leal et al., 1986; Ferreira et al., 1992; Duval et al., 1997),
improved progressively our knowledge of the wild and cultivated forms of the plant, collected them for the morphological,
biochemical, and molecular studies reviewed in the present paper, and gave substance to the hypothesis of a northern origin of
varieties comosus and erectifolius of A. comosus (Leal and Coppens d'Eeckenbrugge, 1996; Coppens d'Eeckenbrugge et al., 1997;
Coppens d'Eeckenbrugge and Leal, 2003; Duval et al., 2003).
The three pineapple domesticates have different stories of domestication, in relation to their different utilizations by man and
regions of origin, so we shall consider them successively.
Ananas comosus var. comosus
The combination of morphological, biochemical and genetic data (Duval et al., 1997, 2001, 2003) clearly point to an
East-Guianese origin of A. comosus var. comosus. Indeed, this area is home to its wild ancestor, A. comosus var. ananassoides.
The greatest phenotypic and genetic diversity, including primitive cultivars and intermediate wild phenotypes that could be used as
a basis for domestication, or that could enrich the primitive cultivated gene pool through introgression, can be found in this region.
A very plausible hypothesis is that such materials were collected on "rock savannahs", sand dunes, and similar places where they
thrive, and planted in home gardens and fields. Nowadays, inhabitants of the Guianese forests, and even creolized newcomers, still
collect materials from the wild to incorporate them in their cultivated plots and gardens. This explains why some clones are found
both under cultivation and in the wild, in patches of secondary vegetation, marking likely sites of ancient cultivation. Such
practices constitute a basis for a process involving "domestication cycles". In these cycles, pineapples are sampled in the wild, put
in cultivation, semi-abandoned, re-sampled for cultivation, etc., with possible selection at each step. Indeed, fields and home
gardens are never completely abandoned and forgotten, as they are located near pathways and remain useful, for example for
picking tree fruits that come well after first crops or hunting animals attracted by the fruits (Vélez, 1998). They also serve as
stocks of useful planting materials. The most interesting genotypes are thus progressively concentrated, in a process that has been
described for Amazonian fruits (Gnecco, 2003; Miller and Nair, 2006). In the long term, sexual reproduction can contribute to the
exploitable diversity by the creation of new clones, some of which can be more attractive for man. On the other hand, wild types
may be more highly fertile than semi-domesticated materials (Coppens d'Eeckenbrugge et al., 1993), so wild genes are probably
transmitted more efficiently by sexual reproduction, reducing the effects of selection. Their robustness may also be an advantage
for a safer harvest, so growers may want to maintain them among the diversity of their clones in a context of subsistence
production . The result is the multiclonal production system still observed in the Guianas, which maintains an equilibrium
between genotypes at very different stages of domestication. This is not a problem for a grower more interested in diversity and
safety than in productivity but slows down further genetic improvement and full domestication.
The relatively slow pace of domestication in other species has also been attributed to the coexistence of genotypes at different
stages of domestication (e.g., Otero-Arnaíz et al., 2005). W ild relatives of domesticates can even behave as weeds in the crop
(Papa and Gepts, 2004), as is the case of teosinte in Mexican maize plots (W ilkes, 1972) or wild sorghum (Dogget and Majisu,
1968) in African fields, contaminating seed materials through pollen-mediated geneflow, and diversifying the cultivated genepool
while braking the evolution to more extreme forms of the crop. Such limitations to genetic improvement obviously disappear when
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Newsletter of the Pineapple Working Group, International Society for Horticultural Science
the crop is grown in the absence of its wild relatives (Galinat, 1974). In the case of pineapple, this probably occurred in the
western Amazon (along the upper Amazon, close to the triple frontier between Colombia, Peru and Brazil, and along the lower
Rio Negro), where we can observe a high diversity of cultivars and the absence of wild types (Bello and Julca, 1993; Duval et al.,
1997). There the prevalence of large seasonally-flooded areas and the rarity of elevated sites with good drainage, such as rock
savannahs and sand dunes, seem to constitute a natural barrier against the extension of the Guianese types of A. comosus var.
ananassoides. Once established in this area, the cultivated pineapple could evolve and diversify in completely artificial conditions,
in a dynamic process combining sexual recombination restricted to domesticated germplasm, somatic mutations and clonal
selection. Human societies peopling western Amazonia may have reinforced the domestication process through particular
horticultural skills. Indeed, this area is also an important centre of domestication and diversification for many other fruits
(Clement, 1989, 1999). Peoples like the Tikunas and the Huitotos still value and maintain a wide diversity of pineapple cultivars
and other fruits. In the course of our collecting trips (Duval et al., 1997), we observed as many as twelve cultivars in a small plot
maintained by a single Tikuna family. Schultes (1991) gives similar numbers for the pineapple cultivars named by the Huitotos.
The species is culturally very important for peoples of the area. For example, the Yukunas celebrate nine fruit festivals yearly, five
of them being pineapple festivals (Jacopin, 1988). Cristancho (2001; cited in Cristancho and Vining, 2004) ranked pineapple
among the three primary Culturally Defined Keystone Species of the Letuama people (C.K.S are "species whose existence and
symbolic value are essential to the stability of a cultural group over time"), along with tobacco and coca.
The existence of two centres for the diversification of A. comosus var. comosus, a primary one in the Guianas, with
diversification involving clones at different stages of domestication, and a secondary one in the western Amazon/Andean foothills,
with recombination between large-fruited cultivars, is also suggested by the distribution of particular leaf margin types. Thus,
genotypes that present a partial suppression of spines are particularly frequent in the Guianas. The most famous such cultivar is
'Smooth Cayenne' itself, the most widely distributed pineapple cultivar, which most commonly presents a few spines at the leaf tip.
This trait is under the control of the S gene, and the allele for the partial suppression of spines is dominant. In the western Amazon
and in the Andes (from Peru to Colombia), leaf smoothness is determined by another gene, named P by Collins (1960). The
dominant allele determines the "piping" trait, where the lower epidermis is folded over the leaf edge, and all spines, except for the
terminal one, are suppressed. The existence of homozygotes for the "piping" gene (Cabral et al., 1997) indicates sexual
recombination among cultivars within this western pool.
Because of very poor conservation conditions in the rain forest and the lack of archaeobotanical research in the Amazon, no
ancient pineapple remains have been found in the two centres of diversification/domestication. Pineapple remains have only been
conserved under arid conditions, and identified in archaeological layers dated from 3,200 to 2,800 BP on the Peruvian Coast
(Pearsall 1992), while seeds and bracts were found in coprolites from the Tehuacan Valley caves dated between 2,200 and 1,300
BP (Callen, 1967). The application of historical linguistics to the names of the pineapple in Ancient Mesoamerica
(glottochronology) also gives us a minimal estimation of the antiquity of pineapple domestication. Consistently,
glottochronological data indicate that the crop was highly significant to Mesoamerican people more than 2,500 years BP (Brown,
2010). Thus, domesticated pineapple was traded and adopted as an important fruit crop at the continental scale more than 3,000
years BP. Such an early extension of its cultivation area implies the preliminary development of cultivars specifically adapted to
the high latitudes and/or altitudes of Peru and Mexico. Indeed, highland cultivars from Andean countries show specific adaptations
and tend to perform poorly in lowland conditions, presenting lower sugar, acidity and firmness, as well as frequent fruit lodging
and deformation. Given the rarity of reproduction through seeds in A. comosus var. comosus, the development of environmentally
specialized cultivars was necessarily a long and slow process, taking place in situ, after the arrival of the domesticated pineapple
in these particular environments. In conclusion, a likely time frame for the divergence between wild and cultivated pineapple lies
between 6,000 and 10,000 years BP, which is consistent with current hypotheses for other major American crops.
Ananas comosus var. erectifolius
A. comosus var erectifolius is morphologically very similar to the variety ananassoides, except for the smooth character of the
leaf. Its very high genetic diversity, its scattering in the phenetic and phylogenetic trees and its proximity to various ananassoides
genotypes, generally from the same origins, suggests that it may have been domesticated several times. According to its
distribution in northern Amazonia and the Antilles, these convergent domestication events quite certainly took place in the Guiana
shield. If we take the reduction in sexual reproduction capacity as an indicator of the antiquity of evolution under domestication,
the highly variable fertility in var. erectifolius appears consistent with multiple domestication and introgression events, as some
clones have partially lost their flowering capacity, whereas others show an abundant production of viable pollen and ovules
(Coppens d'Eeckenbrugge et al., 1997; Duval et al., 2003).
Ananas comosus var. bracteatus
The case of this botanical variety is much simpler, as it very probably corresponds to two particular clones from southern
South America. The most common one (A. bracteatus sensu Smith & Downs) shares its chloroplast genome and most of its
nuclear genome with A. comosus var. ananassoides, and the remaining part of its nuclear genome with A. macrodontes, which
indicates a hybrid origin. The second one presents an even closer affinity with A. macrodontes, as, in the study of Duval et al.
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Newsletter of the Pineapple Working Group, International Society for Horticultural Science
(2003), their chloroplast genomes are only differentiated by one mutation. The two clones have been exploited by man for
multiple purposes, but they were probably more important for fibre, as was their relative A. macrodontes, in the absence of A.
comosus var. erectifolius in the region, than for their poor quality fruits. Their current use as fences is certainly recent, as it was
only justified by the introduction of cattle raising and the development of private property on farmlands.
Beyond the pineapple case
The most obvious interest of the authors and readers of this paper lies in the pineapple, its development as a crop and the
historical and genetic processes that shaped it. However, notwithstanding this sharp interest in the crop, we cannot isolate the
subject from wider contexts. As a fruit crop, pineapple occupies a very special position, because it is one of the most prominent
tropical fruits and the first Amazonian fruit of local and global importance. And it has been so for millennia. Among the fruits of
comparable importance, we can mention avocado (Persea americana Mill.) and papaya (Carica papaya L.), both of
Mesoamerican origin.
The comparison can be extended to their history. All three species are ancient domesticates. Archaeobotanical remains attest
to the antiquity of avocado domestication in Mesoamerica, beginning around 10,000 BP (Galindo-Tovar et al., 2008). For papaya,
we must envisage that management by man is at least as ancient as early lowland agriculture in Mesoamerica, i.e., again
somewhere around 10,000 BP (Piperno, 2006). Indeed, papaya is a very early colonizer of any open space in the forest, and it has
very particular and obvious chemical properties, so early farmers or proto-farmers clearing land for cultivation must have been
very familiar with the plant and the fruit. The three species also provide striking examples of the early diffusion of the most
interesting domesticates across all of tropical America, similar to utilitarian crops, such as cotton and bottle gourd, and the staples,
such as maize and a few important root crops. At the time of Conquest, papaya was present in all of tropical America (Patiño,
2002), where it was introduced in the form of cultivars, as can be easily deduced from the observation of feral papaya populations
in South America, whose fruits differ markedly from the truly wild papayas of Central America. Archaeobotanical remains in
Peru, dated 2800-3200 BP (Pearsall, 1992), confirm the antiquity of its presence in South America. Avocado cultivars may have
diffused to South America as early as 10,000BP, according to seeds found in Colombian archaeological sites (Gnecco, 2003).
Finally, the remarkable diffusion of the three fruits resumed immediately after the contact, as they were successfully transferred to
most tropical regions of the world in less than two centuries. In particular, the pineapple had become a pantropical fruit crop
before the end of the 16th century (Coppens d'Eeckenbrugge et al., 1997). This underlines the clear relation between the qualities
of these plants and their universal acceptance.
Other fruit crops have had more limited prehistoric diffusion, either for ecological reasons, as the peach palm (Bactris
gasipaes Kunth) that remained confined to the humid lowlands, for more limited attractiveness, or because they were domesticated
much later or incipiently. However, all of them merit consideration for the study of plant domestication in Amazonia. Indeed,
American agriculture is very obviously characterized by its great diversity of species, and the high proportion of fruit species,
whose importance in the Amazonian diet traces back as far as the late Pleistocene, more than 10,000 BP, i.e., the same age as the
many fruit remains of the Pedra Pintada archaeological site (Roosevelt et al., 1997). The review of Miller and Nair (2006) shows
the link between this very particular blend of agriculture and agroforestry, based on a great variety of cultivated plants, including
fruit trees, and an autochthonous cultural development, allowing the establishment of large populations with complex societies,
and trade networks so highly efficient that they could reach the Pacific coast beyond the Andes, and even Mesoamerica.
As stated by Zeder (2006), the difference between domestication and other mutualisms essentially lies in the cultural
component and intentionality. If we want to understand how agriculture developed and plant domestication proceeded in the
Americas, and more particularly in Amazonia, we have to keep in mind the close interaction between biodiversity and traditional
knowledge that are integrated and indivisible. As underlined by Vélez (1998), while the viewpoint of western culture (and
science) "is a compartmentalized vision in which a given element can be analysed and defined without considering its relationship
to the whole," indigenous cultures view the same elements "as an integrated whole, considered part of a functional and compact
universe in which the parts are not conceived of separately."
Accordingly, an exaggerated focus on grain and starch plant domestication cannot lead us to a correct understanding of the
development of Amazonian production systems, and it is time to reconsider its major fruit component as much more than "the
cherry on the cake." This means reorienting research efforts, with deeper studies of present-day agrosystems and the distribution
of genetic diversity, and more attention to microbotanical remains of a wider diversity of species in archaeological sites. The
potential benefits of such studies are well worth the efforts, in terms of genetic resources and agricultural/agroforestry
development, as well as archaeological progress. A wider diversity of plants can tell us much more about the people that relied on
them, as exemplified by the study of Brown (2010), who first extracted linguistic information on a group of Mesoamerican plant
species and then reverted the process, characterizing the ecology of ancient peoples' homelands from the requirements of the array
of species they exploited. In this study, less common species allowed a greater resolution in time and space, than more widespread
ones, such as maize and beans.
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Newsletter of the Pineapple Working Group, International Society for Horticultural Science
Acknowledgements
The authors are much indebted to Duane Bartholomew, University of Hawaii, and Charles R. Clement, Instituto Nacional de
Pesquisas da Amazônia, for their valuable contribution through scientific discussion and editing.
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Hornborg, A. 2005. Ethnogenesis, regional integration, and ecology in prehistoric Amazonia. Toward a system perspective. Curr. Anthro. 46:
589-620.
Hawkes, J.G. 1998. Back to Vavilov: why were plants domesticated in some areas and not in others? In : Damania, A.B., J. Valkoun, G. Willcox
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Jacopin, P.Y. 1988. Anthropological dialectics: Yukuna ritual as defensive strategy. Sweiz. Amer. Ges. Bull. 52: 35-46.
Kislev, M.E., Hartmann, A., Bar-Yose, O. 2006. Early Domesticated Fig in the Jordan Valley. Science 312 : 1372-4.
Leal, F., Amaya, L. 1991. The curagua (Ananas lucidus, Bromeliaceae) crop in Venezuela. Econ. Bot. 45: 216-24.
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Leal, F., Antoni, M.G. 1981. Especies del género Ananas: origen y distribución geográfica. Rev Fac Agron, Universidad Central de Venezuela,
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Miller, R.P., Nair, P.K.R. 2006. Indigenous agroforestry systems in Amazonia: from prehistory to today. Agroforest. Syst. 66:151-164
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Wilkes, H.G. 1972. Maize and its wild relatives. Science 177: 1071-7.
Woods, W.I., Teixeira, W.G., Lehmann, J., Steiner, C., WinklerPrins, A., Rebellato, Lilian (eds.). 2009. Amazonian dark earths: Wim
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Zeder, M.A. 2006. Central questions in the domestication of plants and animals. Evol. Anthropol. 15:105-117.—
News from the Philippines
Philippines President Witnesses Signing of $500-M Contract Between the Government and
Libby's Fruits
On Saturday, June 28, 2008, NTC Marketing, a W illiamsville USA-based company that imports tropical fruit sold under the
Libby’s name, signed a deal to buy pineapples from the Philippines. Trade Secretary and National Development (NDC) Chair
Peter Favila signed in behalf of the Philippine Government and NTC Marketing Chairman Michael DeRose signed for Libby's
Fruits. The signing was witnessed by President Gloria Macapagal Arroyo, who was visiting the United States and promoting the
Philippines as a place for U. S. businesses to invest.
NTC Marketing imports millions of cans of fruit each year. It is the North American licensee of Libby’s, the brand under
which the products are sold. According to DeRose, NTC’s chairman, a processing plant is being built in the Philippines and
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Newsletter of the Pineapple Working Group, International Society for Horticultural Science
should be finished by the end of the year. It was reported that the company will have an ownership stake in the new plant and
pineapple fields to supply the cannery have already been planted. Reportedly, the deal will benefit about 5,000 farming families in
the Philippines. NTC also believed the company would benefit, expecting the price would be less than fruit from Thailand,
although the company expected to continue to import fruit from Thailand as well.
Ed. Note: The above information was obtained from two different web pages, one of which is no longer available.—
News from Sri Lanka
Antagonistic effect of Trichoderma harzianum on Thielaviopsis paradoxa - the Pineapple
Black Rot Pathogen
W ijesinghe,C.1 , W ilson W ijeratnam, R.S.1 , Samarasekara, J.K.R.R.2 and W ijesundera, R.L.C.3
Post Harvest Technology Laboratory and 2 Herbal Technology, Laboratory,Industrial Technology Institute, 363, Bauddhaloka
Mawatha, Colombo 07, Sri Lanka; 3 Department of Plant Science, University of Colombo, Colombo 03, Sri Lanka. E-mail:
shanthi@iti.lk
1
Abstract
A local isolate of Trichderma harzianum was tested against Thielaviopsis paradoxa (teleomorph = Ceratocystis paradoxa)
isolated from a diseased pineapple and established as an antagonist of the pathogen. Assays for antibiosis were conducted using
culture filtrates of the antagonist grown in waste residue from beer manufacture (BR) as the highest number of colony forming
units (cfu) of 2.8x106 /ml, and the highest biomass of 7.425g/250ml of BR were observed in this medium 144 hours after
inoculation. W hile the culture filtrates had no inhibitory effect, the unfiltered broth of T. harzianumP3 showed total inhibition.
The fungicidal effect of the antagonist was confirmed by the presence of coiling structures around the pathogen. Growth of T.
harzianumP3 at 28 EC on solid media was slow - 107 cfu 2 weeks after inoculation and 1.4 x 10 9 cfu at 6 weeks in white rice
medium. Finger millet, red rice and rice husk media were also tested. The effect of T. harzianumP3 treatment to
pathogen-inoculated soil was tested. Boiling tubes sterilized with 20 g soil were inoculated with the pathogen and incubated at 28
EC for 7 days. Inoculated soils were checked for presence of 104 ,105 and 106 cfu/mL of the pathogen prior to addition of the test
formulation of T. harzianumP3. Inoculated soils were checked weekly for surviving cfu/mL of the pathogen over 10 weeks. The
pathogen reached concentrations below disease causing levels (103 cfu/mL) at 6 - 10 weeks. This study indicates the possible use
of T. harzianumP3 as a bio-control agent to control black rot disease of pineapple and BR as a potential medium for mass
propagation of T. harzianumP3 in Sri Lanka. Further studies on the efficacy and shelf-life of the formulation are required.
Introduction
The black rot pathogen Thielaviopsis paradoxa (teleomorph = Ceratocystis paradoxa ) is present in all pineapple growing
countries. The pathogen is capable of causing the great economic loss due to rot of pineapples held in storage. In Sri Lanka the
disease is problematic when fruits are harvested in rainy weather and stored prior to shipment. The disease is controlled by low
temperature storage and chemical treatments (Reys et al., 2004). However alternate means of control are needed due to consumer
resistance to chemicals. Trichoderma harzianum has been observed to be antagonistic to several plant pathogens (Perello et al.,
2003; Singh et al., 2007). A local isolate of this organism T. harzianumP3 (TP3) was tested for antagonism against T. paradoxa.
The fungicidal effect of the antagonist was confirmed by the presence of coiling structures around the pathogen. Selected liquid
and solid media were tested for determining presence of antibiosis and developing a formulation for conducting in vitro soil assays
to observe the efficacy of TP3 in controlling black rot. Soils inoculated with T. paradoxa were examined over 10 weeks for
survival (cfu/mL) of the pathogen.
M aterials and methods
Isolation of T. harzianum - An isolate from a pineapple plantation in Sri Lanka was purified on PDA via single spore method
and given the culture number T. harzianumP3 (THP3). Seven day old cultures of THP3 on PDA were used to prepare the spore
suspensions. Sterile water (10ml) was added to plates, the culture surface agitated, and the resulting suspension filtered thro ugh
sterile cotton wool. The spore concentration of the suspension was adjusted to 1 x 10 7spores mL -1.
T. paradoxa - The pathogen, was isolated form the stem end of a naturally infected pineapple. Pure cultures were maintained on
PDA at 28 ºC, and pathogenicity confirmed via Koch's postulates (Reyes et at 2004). Spore suspensions of the pathogen were
prepared as described above.
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Newsletter of the Pineapple Working Group, International Society for Horticultural Science
Bio assay 1 - Four sterile PDA plates were used for the assay. Four wells of diameter 4 mm were cut equidistant from each other.
Each well was filled with 75 µl of the 107 spore suspension of THP3. A mycelial disc of T. paradoxa obtained from a 7 day culture
was placed in the centre of each plate prior to incubation at 28 ºC for 8 days.
Bio assay 2 - The pathogen was grown in liquid medium to investigate possible antibiosis. Flasks containing 100 mL of the waste
residue from beer manufacture (BR) medium, acidified with tartaric acid, were sterilized and inoculated with 1mL of 1x10 7 TP3
spore suspension for incubation at 28 ºC in a 120 rpm rotary shaker for 7 days. W ells were cut on PDA plates as described
previously. W ells of five plates were filled with 75 µl of culture filtrate (0.45 µm sterile membrane filter, Sartorius AG, 3400
Göttingen, SM 16510), while wells of a second set of five plates were filled with 75 µl of sterilized un-inoculated BR broth. Wells
of a third set of five plates were filled with 75µl of an unfiltered 7 day culture broth. All plates were centrally inoculated with a 4
mm mycelial disc taken from the periphery of a 7 day culture of T. paradoxa and incubated at 28 ºC for 7 days for daily
observation. Slide cultures were prepared using the method described by Sivakumar et al., 2000.
TP3 growth in Liquid and Solid media - Potato Dextrose Broth, (PDB) and Czapekx Dox medium were prepared as in the
Difco manual (257-258, 691-692). Brewery waste (BR) was prepared by dissolving 20 g of BR (Lion Breweries, Biyagama, Sri
Lanka) and 20 g of glucose in 1000 mL of water. Autoclaved tartaric acid (1.6 mL) was added to sterilized media (250ml) to
prevent bacterial growth. Each media containing flask was inoculated with 2.5 mL of 1x10 7 TP3 spore suspension, and incubated
at 28 EC in a 120 rpm rotary shaker. The cultures were harvested between 48 and 360 h after inoculation by aseptic filtration
(W hatmann no: 125 filter paper). The residual mycelium was dried at 80 EC to a constant weight. The culture filtrate (1 mL) was
serially diluted, and plated for recording cfu/mL from each dilution for each medium after incubation at 28 EC (Singh et al.,
2007).The experiment was carried out in triplicate.
Red rice, white rice, paddy husk and finger millet were used as solid media. Each medium (10g) was soaked over night in
10mL water and cooked with 0.5 mL of 10% sunflower oil, cooled, placed in polypropylene bags and sterilized. Soil dilution plate
technique was used to record cfu/g from each of four replicate bags inoculated with 10 7 spore/mL of TP3 for each medium, after 2
weeks incubation at 28 ± 2ºC.
Bio Assay 3 - The in vitro efficacy of TP3 was tested using boiling tubes sterilized with 20 g of soil. Four replicate tubes were
inoculated with the pathogen at disease causing concentrations of 104 ,105 ,106 spores per mL and incubated at 28 EC for 7 days.
The cfu/g of the pathogen in inoculated soils was checked prior to addition of 10 mL of a formulation (authors unpublished data)
of 107 spores per mL of TP3. Inoculated soil samples were incubated at 28 EC. Efficacy of the formulation was checked by
recording the number of cfu/g of pathogen each week over a period of 8 weeks.
Statistical Analysis - All results were statistically analyzed using the Statistical Analysis System (SAS) computer package Version 6. Mean values were compared by the least significant difference test (LSD) at 5% level of confidence.
Results
The dominance of the antagonist was evident 3 days after inoculation in bio-assay 1. All plates were covered by TP3 on day 7
and it was not possible to re-isolate the pathogen from the assay plates at this stage. Growth of TP3 at 28 EC on solid media was
slow with best results of 107 cfu /g, 2 weeks after inoculation and 1.4 x 109 cfu/g at 6 weeks in white rice medium. In liquid media,
the highest number of 2.8x106 cfu/mL and the highest biomass of 7.425g/250ml of TP3 was observed in BR medium 144 hours
after inoculation (Figures 1 and 2). Inhibition of T. paradoxa was observed only in the unfiltered crude broth of TP3, with total
inhibition observed 5 days after inoculation. Results suggest that extra-cellular metabolites do not play a role in growth inhibition
of the pathogen in this case. The fungicidal effect of the antagonist was confirmed by the presence of coiling structures around the
pathogen using slide culture technique. The pathogen reached concentrations below disease causing levels (103 cfu/mL) at 6-10
weeks (Figure 1). This study indicates the possible use of T. harzianumP3 as a bio-control agent to control black rot disease of
pineapple and BR as a potential medium for mass propagation of T. harzianumP3 in Sri Lanka. Studies on the efficacy and
shelf-life of the formulation are in progress and are to be followed by field trials.
References
Reyes, M.E.Q., Rohrbach, K.G., Paul, R.E., 2004. Microbial antagonists control post harvest black rot of pineapple fruit. Post harvest biology
and Technology 33, 193-203.
Perello, A., Monaco, C., Simon, M.R., 2003. Biocontrol efficacy of Trichoderma isolates for tan spot of wheat in Argentina. Crop Protection 22,
1099-1106.
Singh, H.B., Shishir, S., Singh, A., 2007. Effect of substrates on growth and shelf life of Trichoderma harzianum and its use in biocontrol of
diseases. Bio-resources Technology 98, 470-473.
Sivakumar, D., Wilson Wijeratnam R.S., R.L.C. Wijesundera, F.M.T. Marikar, M. Abeysekere (2000) Antagonistic effect of Trichoderma
harzianum on Post harvest pathogens of rambutan Nephilium lappaceum. Phytoparasitica, 28(3)240- 247.
29
Newsletter of the Pineapple Working Group, International Society for Horticultural Science
Figure 1. Growth of T. harzianum P3 on selected solid media at 28 °C.
Figure 2. T.harzianum P3 cfu/m l in liquid m edia .
30
Newsletter of the Pineapple Working Group, International Society for Horticultural Science
Figure 3. Effect of T.harzianum P3 formulation on 104, 105, and106 spore/mL of T.paradoxa
in-vitro.
—
News from Taiwan
Invited short note
Forced Flowering of Pineapple (Ananas comosus cv. Tainon 17) With Calcium Carbide Plus
Activated Charcoal and By Ice Cold Stress
Chin Ho Lin and Subbiyan Maruthasalam. Department of Life Sciences, National Chung Hsing University, Taichung 402, Taiwan,
ROC, chinho@dragon.nechu.edu.tw
Increasing the efficacy of calcium carbide
In pineapple cultivation, synchronization of flowering is an important agronomic practice that is severely disrupted by
induction of natural flowering (natural induction) associated with the gradual decline in night temperature and shortened day
lengths during the winter months. This problem is more serious in sub-tropical countries like Taiwan (latitude 22° to 26°N). In
subtropical environments it is assumed that natural induction in pineapple is a physiological response to ethylene production in the
shoot apical meristem due to low night temperature. Recent studies showing inhibition of natural induction with the ethylene
biosynthesis inhibitor aviglycine (W ang et al., 2005; W ang et al., 2007), has proved this assumption. Once natural induction is
prevented, flowering can be restored later at convenience through forcing with ethylene or calcium carbide (CaC 2). In Taiwan,
CaC 2 remains primary choice for forcing of pineapple because of low cost and ease of application. A grower with the sole of aim
of obtaining synchronized forcing would make two or more CaC 2 treatments, thus resulting in increased cost and reduced profit.
In vitro studies showed that the acetylene released from CaC 2 upon the addition of water was efficiently absorbed by activated
charcoal and significant acetylene release could be detected even after 24 h. W e then explored this property under field conditions
by supplementing 0.5-1.0% CaC 2 with 0.5-2.0% activated charcoal. Van de Poel et al. (2009) reported that ethylene absorption
needed at least 5% activated charcoal. This may be explained by the possible variation in the absorptive capacity of activated
charcoal towards acetylene and ethylene. About 50 ml of aqueous solution containing CaC 2 with or without charcoal was applied
into the leaf rosette of 11-month-old ‘Tainon17’ (‘Smooth Cayenne’ × ‘Queen’) plants during the 2007-2008 season.
Supplementation of 1.0% CaC 2 with 0.5% activated charcoal improved the forcing effectiveness by up to 40%, while 1.0-2.0%
activated charcoal drastically reduced the forcing percentage. Based on these results, taken together, we could assume that 1.0%
CaC 2 containing 0.5% activated charcoal applied once or 0.5% CaC 2 containing 0.5% activated charcoal applied twice would
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Newsletter of the Pineapple Working Group, International Society for Horticultural Science
force pineapple. Thus, activated charcoal provides the flexibility of reducing the number or concentration of CaC 2, or both, to
force pineapple and reduce cost.
Forcing with ice and ice water
Though ethylene is permitted for forcing of organically grown pineapple in many countries, it still is considered unnatural by
some consumers, thus making it desirable to develop a more acceptable organic method of forcing in pineapple. Organic forcing
can be defined as forcing of pineapple with any agent that can elicit the endogenous production of ethylene in shoot apical tissues
and initiate reproductive development without the exogenous application of ethylene or related compounds. In our laboratory, gas
chromatographic analyses showed that treating shoot apical tissues of pineapple with ice for one hour stimulated a two-fold higher
ethylene production than comparable apical tissues similarly treated with water at 25 °C. This prompted us to evaluate the effect of
ice treatments under field conditions. ‘Tainon17’, a cultivar showing high sensitivity to natural induction was selected for the field
tests. Ice treatments were applied to 11-month-old plants (3.5-4 kg mass) in the third or fourth week of October during 2006-2007
and 2007-2008 seasons. Plants were treated 1-4 times with from 500 g to 2 kg of ice crystals or 500 ml of ice water at 24 h
intervals. Inflorescence emergence (budding) in the leaf whorl was counted and expressed as a percentage. Plants treated with
CaC 2 (2X at a 48 h interval) , ethephon (2X at a 48 h interval) and water (25°C) served as controls.
In the 2006-2007 season, budding of plants treated once or twice with ice or ice water was not observed until the third week
of February. Nearly 100% budding was observed for plants treated with CaC 2 (2X-48 h interval) by the second week of December.
Though ice or ice water treated plants are not induced to flower with the relative efficacy of CaC 2, bud emergence from coldstressed plants was 1-2 weeks earlier than those treated with water (25°C), indicating that cold treatment altered the sensitivity of
plants to natural induction. In the 2007-2008 season, plants treated 3 or 4 times with ice or ice water were induced to flower with
an effectiveness almost equal to that of CaC 2 and ethephon. However, bud emergence from the cold-treated plants was delayed by
2-3 weeks. These experimental results indicate that under field conditions, 1-2 ice treatments were not sufficient to force flowering
of ‘Tainon 17', but 3-4 ice treatments readily induced flowering in ‘Tainon17’. Interestingly, four applications of 500 ml of ice
water resulted in better forcing than did four applications of 500 g of ice and was equal to the results obtained on treating plants
four times with 2.0 kg of ice. It is difficult to explain this scenario without data on the temporal fluctuations in the ACC synthase
activity after ice-cold treatments. As expected, maturity and harvesting of fruits from ice treated plants was delayed by 2-3 weeks
compared to CaC 2 and ethephon and in most treatments harvesting was completed in 3-4 harvest passes over 7-10 days.
Morphology of fruits from ice treated plants showed no discernable variation from that of chemically-forced fruits, however, the
qualitative parameters were not studied.
It is assumed that the high sensitivity of ‘Tainon17’ to natural induction is what accounts for the relative ease in forcing this
cultivar with ice treatments in Taiwan in October, i.e. 2-3 weeks prior to natural induction. Soler et al. (2006) reported that cold
water (5°C) treatment was more effective on those cultivars that are sensitive to natural induction. Forcing efficiency heavily
depends on the cultivar sensitivity and prevailing weather conditions (Bartholomew et al., 2003) and, therefore, the effect of ice
treatment on inflorescence emergence should be tested, (i) on cultivars differing in their sensitivity to natural flowering, and (ii)
under varying climatic conditions and production seasons, before incorporating the method into the regular farming schedule.
Acknowledgement
The authors thank D. P. Bartholomew, University of Hawaii at Manoa, Honolulu, USA, for his valuable comments on the
experimental results and critical reading of this note.
References
Bartholomew, D.P., Malezieux, E., Sanewski, G.M., Sinclair, E., 2003. Inflorescence, and fruit development and yield. In: Bartholomew, D.P.,
Paull, R., Rohrbach, K.G. (Eds.), The pineapple: Botany, production and uses. CABI Publishing, Wallingford, U.K, pp. 167-202.
Kuan, C.S., Yu, C.W., Lin, M.L., Hsu, H.T., Bartholomew, D.P., Lin, C.H., 2005. Foliar application of aviglycine reduces natural flowering in
pineapple. HortScience 40, 123-126.
Soler, A., Teisson, C., Dole, B., Marie Alphonsine, P.A., 2006. Forcing in pineapple: what is new? In: Bartholomew, D.P. (Eds.), Pineapple
News 13, 27–30; http://www.ishs-horticulture.org/workinggroups/pineapple/.
Van de Poel, B., De Proft, J.C.M.P., 2009. Determination of pineapple (Ananas comosus, MD-2 hybrid cultivar) plant maturity, the efficiency
of flowering induction agents and the use of activated carbon. Sci. Hort. 120, 58-63.
Wang, R.H., Hsu, Y.M., Bartholomew, D.P., Maruthasalam, S., Lin, C.H., 2007. Delaying Natural flowering in pineapple through foliar
application of aviglycine, an inhibitor of ethylene biosynthesis. Hortscience 42, 1188–1191.—
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Newsletter of the Pineapple Working Group, International Society for Horticultural Science
News From the United States (Hawaii)
Phytophthora Heart Rot Control
B. Sipes1 G. Taniguchi1 and T. Radovich2 . Depts. 1 Plant & Environmental Protection Sciences and 2Tropical Plant & Soil Science,
Univ. Hawaii at Manoa, Honolulu, HI 96822. E-mail: sipes@hawaii.edu
In a preliminary field test, we evaluated a vermicompost tea drench and a BTH (acibenzolar-s-methyl) dip as compared to an
Aliette dip and an untreated control on the incidence of Phytophthora heart rot of pineapple. Plots were prepared by running an
irrigation tube down the center of a bed and covering the bed with plastic mulch. The treatments were arranged in a RCBD with
plots 1 bed (1 m) wide by 1.5 m long. Blocks consisted of one 4.5 m row. The four treatments were replicated 10 times in plots of
5 plants. Crowns of hybrid 73-50 were dipped in water, Aliette (300 g/100 l), or BTH (100 ppm) and then planted. For the
vermicompost tea, water-dipped crowns were planted and then drenched with 500 ml tea/crown (1.8 mt of compost in 23,333 L
ha-1 water). Crowns in the other treatments were drenched with 500 ml water. The plots were irrigated for 48 hours after planting
and then again weekly for the next 12 weeks. Crown rot was recorded 3, 6, 9, and 12 weeks after planting.
More crowns (38%) were lost in plots receiving the compost tea treatment than in any other treatment (Figures 1). Only one
Figure 1. Number of dead and dying pineapple crowns under different dip or drench
treatments.
crown was lost in the Aliette treatment after 12 weeks. W e suspect this dead plant was actually a 73-114 hybrid contaminant based
upon its leaf characteristics. Sixteen percent of the crowns were lost in the untreated plots whereas only 8% of the crowns were
lost in the BTH treatment.
This vermicompost tea appeared to either stimulate or encourage Phytophthora infection. The BTH, a known inducer of
systemic acquired resistance, may have induced resistance to phytophthora in these pineapples as well. Since the hybrid 73-50 is
more tolerant/resistant to phytophthora than 73-114, we are repeating the evaluation using 73-114 in hopes of achieving greater
infection in the untreated plots.
Evaluation of Transgenic Resistance
B. Sipes, C. Nagai, and M.-L. W ang
W e have evaluated 13 lines genetically modified with a cystatine gene to control nematode reproduction in pineapple. In a
group of 17 different pineapple lines (the number of individual plants per line varied), root-knot nematode (Meloidogyne
javanica) infection reduced pineapple growth an average of 43% in 11 of the lines. In the remaining two lines, plants grew more in
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Newsletter of the Pineapple Working Group, International Society for Horticultural Science
the presence of nematodes, however these were generally the smaller plants in the group. All of the plants supported root-knot
nematode reproduction. Two lines had fewer numbers of nematode than the other lines but were not among those lines undamaged
by the nematodes. W e compared nematode reproduction and plant growth of one of the transgenic lines (composed of 350 plants)
to that of a wild type (composed of 235 plants) in the greenhouse. Overall the transgenic plants were 37% smaller and supported
42% more nematodes per plant than the wild type plants. The cystatin levels were probably not high enough in the transgenic line,
even though this line had the highest expression level of all lines based on western blot analysis. The level recorded was much
lower than what has been reported in other transgenic plants like potato.
Table 1. Root-knot nematode (Meloidogyne javanica) reproduction on transgenic and wild type pineapple.
Plant
Nematode
Initial plant
weight (g)
Transgenic
Transgenic
+
5.03
4.97
W ild
W ild
+
5.5
5.34
Final
Root
Shoot
plant
weight
weight (g)
weight (g)
(g)
74.85
66.41
4.74
72.75
65.15
4.55
118.34
119.34
104.15
103.87
7.52
7.94
Dry root
weight (g)
Total
nematode
Nematode/g
dry root
1.45
1.42
3
2366
1661
2.11
2.31
6
1670
722
Production of transgenic pineapple (Ananas cosmos (L.) Merr.) plants via adventitious bud
regeneration (accepted by In Vitro Cellular and Developmental Biology - Plant)
M.-L. W ang, G. Uruu, L. Xiong, X. He, C. Nagai, K. T. Cheah, J. S. Hu, G.-L. Nan, B. S. Sipes, H. J. Atkinson, P. H. Moore, K.
G. Rohrbach, R. E. Paull
A new protocol for the production of transgenic pineapple plants was developed. Adventitious buds were induced directly
from Agrobacterium-infected leaf bases and stem discs of in vitro plants, bypassing the establishment of callus cultures.
Non-chimeric transgenic plants were obtained by multiple subculturing of primary transformants under increasing levels of
selection. A total of 42 independent transgenic lines were produced from two cultivars with two different constructs; one
containing a modified rice cystatin gene (Oc-IÄD86) and the other with the antisense gene to pineapple aminocyclopropane
synthase (ACS). GUS histochemical staining provided the first evidence of the non-chimeric nature of the transformed plants.
Their non-chimeric nature was further demonstrated by PCR analyses of the DNA extracted from individual leaves of a primary
transformed plant and also from multiple plants propagated from a single transformation event. Southern hybridization confirmed
random integration patterns of transgenes in the independent lines. The expression of Oc-IÄD86 gene was detected via RT-PCR at
the mRNA level and translation by protein blot. Agronomic evaluation and bioassays of the transgenic plants will further validate
the utility of this new tool for pineapple improvement.
Effects of ReTain® on Natural Induction of Reproductive Development of MD-2 Pineapple
Duane P. Bartholomew and Gail Uruu. Dept. of Tropical Plant and Soil Science, University of Hawaii, Honolulu, HI 96822.
E-mail: duaneb@hawaii.edu
Natural induction (NI) of reproductive development of pineapple is a serious problem for pineapple growers and particularly
for those growing cultivars highly sensitive to NI. The problem is more severe in regions with cool winter temperatures and
shorter daylengths. In such environments, the incidence of NI can be high, resulting in off-schedule fruiting, a spread harvest peak
and increased variability in ratoon crop fruits due to variation in sucker initiation and development. In Hawaii, NI can occur as
early as the middle of November and it can continue into March.
Aviglycine, an ethylene biosynthesis inhibitor, can control NI in pineapple (Kuan et al., 2005; Lin et al., 2006; W ang et al.,
2007). However, aviglycine is very costly and further studies are needed to reduce costs while maintaining efficacy in areas where
NI is a serious problem. In a 2007-2008 test with aviglycine (Bartholomew and Uruu, 2008), treatments were begun on December
1 and significant NI occurred on or soon after that date, which resulted in less than ideal control of the problem. As a follow-up to
that study, two tests of the efficacy of aviglycine (ReTain® and VBC-30102, a liquid formulation of aviglycine) in controlling NI
of ‘MD-2' (Dole’s MG3) were conducted during the 2008-2009 winter season. The tests were located in fields having ‘MD-2'
plants of similar age and plant size. The treatments (Table 1) were designed to test the efficacy of weekly sprays of 100 or 200 mg
L-1 of aviglycine and of a biweekly spray of 100 mg L -1 of aviglycine in controlling NI. Increasing the concentration to 200 mg L -1
34
Newsletter of the Pineapple Working Group, International Society for Horticultural Science
and halving the volume would double the area that could be covered by one tank of spray solution. Treatments with delayed
starting dates were included to help identify more precisely when NI occurs in fields of ‘MD-2' pineapple in Hawaii. The label for
aviglycine only permits a maximum of 10 applications so delaying the application of aviglycine as long as possible offers the
potential to reduce costs or extend coverage later into the winter months, or both. Treatments were first applied on November 7,
2008 in W aialua field 7 (145 m elevation) and on December 1, 2008 in Brodie field 13 (275 m elevation).
Table 1. Treatments to evaluate the efficacy of ReTain® in controlling natural induction of flowering in MD-2 pineapple during
2008-09.
Treatment
Product
A.I., mg L-1
L acre-1
Applications
Est. No.
Interval, days
T1
Control
----T2
ReTain®
100
946.3*
3-5
14
T3
ReTain®
100
946.3
6 - 10
7
T4
ReTain®
200
473.1
6 - 10
7
T5
VBC-30102
100
946.3
6 - 10
7
T6
VBC-30102
200
473.1
6 - 10
7
T7†
ReTain®
100
946.3
3-7
7
T8†
ReTain®
100
946.3
2-4
7
* Volume of 946.3 L = 250 gallons/acre and 473.1 L = 125 gallons/acre.
†Aviglycine was applied approximately one (T7) or two (T8) weeks later than the other treatments.
Plot size was 4 two-row beds by 10 m and the experimental design was a randomized complete block (RCB) with four
replications. Bud emergence data were only obtained for an average of 137 plants in the two inner beds; the outer two beds served
as buffers between adjacent plots. Because of time constraints, the incidence of NI was not determined in all plots. The tap wateraviglycine solution was applied as a broadcast spray in either one (473.1 L) or two (946. L) passes over the plots using a 0.98 m
(60") boom with 6 TeeJet fan-spray nozzles spaced 20 inches apart and a spray pressure of 30 psi. No surfactant or other adjuvant
was used.
The treatments were usually applied between 7:30 and 9:00 a.m. at the W aialua location and between 9:30 and 11:00 at
the Brodie location. Air temperature data collected in the field made it possible to evaluate the effect, if any, of time of
application and associated temperature changes on the efficacy of aviglycine treatments. To keep the total application time as
short as possible, bulk volumes of the various sprays were mixed in the tank and then all plots for those treatments in each test
were sprayed before another spray solution was mixed. The tank and hose were flushed with water between sprays having
different concentrations or formulations. At the W aialua test site treatments were applied in the order 100 mg L -1 ReTain ® , 200
mg L-1 ReTain, 100 VBC-30102 followed by 200 mg L-1 VBC-30102. To minimize the time spent cleaning the spray tank and
flushing the hose and boom, the last treatment at the W ailua location was the first treatment applied at the Brodie location. The
last treatments at Brodie were always those containing 100 mg L -1 of ReTain ®.
To the extent possible, the dates of NI were estimated using bud counts collected from 10 plants forced with 10 cc of a
solution containing 4% urea and 45 mg a.i. of ethephon on each treatment date at each location. In some of the plots forced during
January and February, it was not possible to distinguish between forced plants and those induced naturally. Bud counts for the
treated plots were converted to percentage NI and the data were arcsine transformed prior to statistical analysis using the GLM
procedure. The means were retransformed for reporting purposes.
The average air temperature range for the dates and times the treatments were applied at the two sites gradually decreased
over time at both locations (Table 2, Table 3). Although average air temperature at the W aialua site was about 1.0 EC (1.8 EF)
warmer than at the Brodie site (data not shown), the temperatures during the time the treatments were applied was warmer at
Brodie than at W aialua because of the later starting time at the Brodie field. Days to budding (approximately 1 cm open heart) of
the forced plants, where they could be determined unambiguously, ranged from 52 to over 70 days at the two locations. Daily heat
units (mean air temperature minus a base or threshold temperature of 12 EC (59 EF); Fleisch and Bartholomew, 1987) were
calculated and summed over days to about 1.0 cm open heart for each location. W here bud emergence could be clearly associated
with forcing, as opposed to NI, the accumulated heat units from forcing to 1.0 cm open heart were quite similar at both locations.
These results confirm the results of Fleisch (1988) who reported that accumulated heat-units is a simple and reliable predictor of
fruit development up to the time of bud emergence.
Control of natural induction (NI) at the W aialua site in early March (Table 4) was excellent when the treatments were
begun before November 24, 2008. Applying aviglycine at biweekly intervals or delaying the treatment start date until November
24 still resulted in a highly significant reduction in NI as compared with the control. Treatment with 100 or 200 mg L -1 aviglycine
at weekly intervals provided excellent control of NI and the liquid formulation of aviglycine was as effective as the powder
formulation. The excellent control of NI achieved with weekly applications of 100 mg L-1 aviglycine suggests that it would be
worthwhile to evaluate the efficacy of lower concentrations of aviglycine in hopes of reducing the cost of product required to
35
Newsletter of the Pineapple Working Group, International Society for Horticultural Science
prevent NI. Biweekly application of aviglycine also appears to offer significant improvement in control of NI and could offer
significant savings in cost of product or double the period during which NI might be controlled.
Table 2. Application date, time and temperature range when aviglycine treatments were made and budding information for plants
forced on the various treatment dates at W ailua field 7, block 4.
Date
Time
Temperature
Budding‡ date, days to budding and cumulative heat units
Date
Days
Heat units
Nov. 7
8:00 - 9:00
No data
Dec. 29
52
Nov. 14
8:00 - 9:00
No data
Jan 5
52
Nov. 24
8:00 - 10:00
23.4 - 25.9
ND*
Dec. 1
7:50 - 9:15
21.0 - 22.2
Jan. 26
56
526 (915)†
Dec. 8
7:35 - 9:15
20.0 - 24.8
Feb. 6
60
550 (1008)
Dec. 15
7:50 - 9:20
21.4 - 24.7
ND
Dec. 22
7:40 - 9:10
18.9 - 21.0
ND
Dec. 29
7:45 - 9:00
18.6 - 23.4
ND
Jan. 5
7:45 - 9:10
19.5 - 23.2
ND
Jan. 12
7:45 - 9:10
16.4 - 20.4
ND
‡Budding stage was 1 cm open heart.
*Not determined because natural induction confounded the observations.
†Units are Celsius or, in parentheses, in Fahrenheit.
Table 3. Application date, time and temperature range when aviglycine treatments were made and budding information for plants
forced on the various treatment dates at Brodie field 13, block 13.
Date
Time
Temperature
Budding‡ date, days to budding and cumulative heat units
Date
Days
Heat units
Dec. 1
10:00 - 12:00
21.6 - 27.3
Feb. 2
63
512 (921)†
Dec. 8
9:45 - 10:45
23.5 - 25.2
Feb 20 est.
74
568 (1070)
Dec. 15
9:50 - 11:20
26.1 - 29.4
ND*
Dec. 22
9:50 - 11:00
21.4 - 23.3
Mar. 2
69
523 (942)
Dec. 29
9:45 - 11:05
24.8 - 27.1
ND
Jan. 5
9:45 - 11:00
22/1 - 26.3
ND
Jan. 12
9:45 - 11:00?
24.4 - 29.4
ND
Jan. 19
7:? - 9:?
13.3 - 19.1
ND
Jan. 26
7:40 - 9:10
15.5 - 19.1
ND
Feb. 2
7:50 - 9:10
18.4 - 20.1
ND
‡Budding stage was 1 cm open heart.
*Not determined because natural induction confounded the observations.
†Units are Celsius or, in parentheses, in Fahrenheit.
Table 4. Effect of aviglycine on percentage of natural induction of reproductive development of MD-2 pineapple in W aialua Field
7, Block 4. Treatments were begun on the indicated starting date in 2008 and all were ended on January 12, 2009. The means,
retransformed after analysis, are based on bud counts for an average of 137 plants per plot made on March 2, 2009.
Treatment
Start
Interval
Replications
Budding, %†
date
days*
Control
Nov.-7
NA
4
53.8 a
ReTain®, 100
Nov.-7
14
4
12.4 b
ReTain®, 100
Nov.-7
7
4
1.2 cd
ReTain®, 200
Nov.-7
7
2
1.6 bcd
VBC-30102, 100
Nov.-7
7
4
0.1 d
VBC-30102, 200
Nov.-7
7
4
1.0 cd
ReTain®, 100
Nov.-14
7
2
1.0 cd
ReTain®, 100
Nov.-24
7
2
8.4 bc
*ReTain spray interval in days.
†Means followed by the same letter are not significantly different from each other, P = 0.5.
Table 5. Effect of aviglycine on percentage of natural induction of reproductive development of MD-2 pineapple in Brodie Field
13, Block 13. Treatments were begun on the indicated date and all were ended on February 2, 2009. The means, retransformed
after analysis, are based on bud counts made on March 9, 2009.
36
Newsletter of the Pineapple Working Group, International Society for Horticultural Science
Treatment
Start
Interval
Replications
Budding, %†
Budding, %
date
days*
<LC‡
<MC
Control
Dec. 1
NA
4
43.5 a
30.9 ab
ReTain®, 100
Dec. 1
14
4
17.1 b
14.8 bc
ReTain®, 100
Dec. 1
7
4
6.3 bc
4.3 cd
ReTain®, 200
Dec. 1
7
3
4.9 bc
2.8 cd
VBC-30102, 100
Dec. 1
7
4
0.7 c
0.5 d
VBC-30102, 200
Dec. 1
7
4
7.0 bc
5.6 cd
ReTain®, 100
Dec. 8
7
4
15.0 b
12.4 bc
ReTain®, 100
Dec. 15
7
4
51.0 a
37.6 a
*ReTain spray interval in days.
†Means followed by the same letter are not significantly different from each other, P = 0.5.
‡The column “Budding <LC” excludes data for plants where the inflorescence was at or more mature than the late cone stage from
the analysis and the column “Budding <MC” similarly excludes data for plants at or older than the mid cone stage of development
because these plants likely began inflorescence development prior to the start of the treatments. Detailed staging was based on the
inflorescence development stages ½" open heart,1" open heart, early cone, mid cone, late cone, early flower, mid flower, late
flower and dry petal. The respective stages occur at approximately weekly intervals in Hawaii.
Delaying application of aviglycine until December 1 can result in increased NI. Since aviglycine treatments at the Brodie
location were not begun until December 1, it was important to be able to distinguish between buds that resulted from NI before the
treatments began from those developing afterwards. This was done by comparing bud development of plants in an adjacent plot
forced on December 1, 2008 with bud development of plants within the test plots. Bud counts in the test plots were made on
March 9, 2009 and buds that were comparable in size to or more developed than those in the plot forced on December 1 were
excluded from the analysis. Because it was difficult to precisely identify the stage of development in both the forced plot and the
test plots, two data sets were created, one where buds more mature than the late cone stage (Rohrbach and Johnson, 2003) of
development were excluded and another where plants with buds more mature than the mid cone stage were excluded. Plant
numbers per plot were reduced to as few as 65 in one plot but for most plots the analysis was based on 100 plants or more. The
results of the two analyses (Table 5) were similar and also were comparable to those obtained for the W aialua block. W e
conclude that application of aviglycine later in the day when temperatures were warmer did not reduce the effectiveness of the
treatments. As was the case for the W aialua test, the least amount of NI occurred in plots that were treated weekly with 100 or
200 mg L-1 of aviglycine. In the <LC analysis, biweekly treatment with aviglycine controlled NI significantly better than no
treatment; however, in the <MC analysis, the control and biweekly treatments were not different from each other. There was a
significant increase in NI when treatments were delayed until mid December. From the 2008-2009 data, for best control of NI in
the Hawaii environment treatments with aviglycine need to be started by late November.
Fruit development was assessed on May 5, 2009 at both the W ailua and Brodie locations. The range of fruit
development spanned several weeks and included fruits that were at the 1.0 cm open heart stage to well beyond the dry petal stage.
Some harvest data will be collected but the wide range of ages and the lack of labor for harvesting will limit the data collected to
the W aialua test.
ReTain® Applied During Early Fruit Development Causes Fruit Deformities
To date, no reports of any detrimental effects of ReTain on fruit development were found. However, in this study,
ReTain sprays inhibited fruitlet development quite significantly. An unusually wide bed interspace separated beds three and four
in replication 3, treatment 8 (R3T8) and replication 4, treatment 3 (R4T3). There was a high percentage of NI in the two rows
bordering the wide interspace. The wide interspace had no effect on ReTain spray coverage of plants in the outside row of bed
three but resulted in little or no ReTain spray coverage of the insdie row of bed four in the two plots. ReTain sprays (100 ppm)
were first applied to R4T3 on December 1 and, when compared to untreated fruit, development of most fruitlets of some fruit was
inhibited (Figure 1). The extent of the growth inhibition/deformity was less in R3T8 and was visible only on the upper portion of
fruits where ReTain sprays were first applied on December 15, 2008 (Figure 2). It appears that the growth inhibition due to
application of ReTain is more pronounced when sprays are applied at about the same time that development associated with NI
begins. Deformities similar to those observed in this test were also seen in plantation blocks that were treated with ReTain
beginning on December 1, 2008.
37
Newsletter of the Pineapple Working Group, International Society for Horticultural Science
Figure 1. Fruits from untreated plants (top row) and fruits from plants treated weekly with 100 ppm ReTain® sprays beginning
on December 1, 2008.
Figure 2. Fruits from untreated plants (top row) and fruits from plants treated weekly with 100 ppm ReTain® sprays beginning
on December 15, 2008.
The percentage of deformed fruits was relatively to very high in all treated plots in the Brodie test (Table 1). The highest
percentage of deformed fruits was in the treatment where ReTain sprays were delayed until December 15. The results indicate that
ReTain treatments must begin before NI occurs, not only to prevent NI but also to avoid ReTain-induced fruit deformity.
Table 6. Effect of ReTain sprays on Percentage deformed fruit in ReTain test.
Treatment
Control
ReTain®, 100 ppm, 14 day interval
ReTain®, 100 ppm, 7 day interval
ReTain, 200 ppm, 7 day interval
VBC-30102, 100 ppm, 7 day interval
VBC-30102, 200 ppm, 7 day interval
ReTain, 100 ppm, 7 day interval beginning Dec. 8
ReTain, 100 ppm, 7 day interval beginning Dec. 15
Deformed fruit, %
0.32
8.42
11.30
19.23
9.86
11.86
19.37
34.46
Acknowledgements
Help with installation of the tests provided by Jenny Antonio and her staff was gratefully appreciated. The weather data was
provided by T. Matsumoto and D. Aoki of the USDA in Hilo, Hawaii.
References
Bartholomew, D.P. and G. Uruu. 2008. Pineapple control of flowering studies. Pineapple News No. 15, pp. 34-38.
Fleisch, H. 1988. Modeling pineapple plant growth and inflorescence development. Ph.D. dissertation. University of Hawaii at Manoa, 195 pp.
Fleisch H., Bartholomew D.P. (1987) Development of a heat unit model of pineapple ('Smooth Cayenne') fruit growth from field data. Fruits
42:709-715.
Kuan C., Yu C., Lin M., Hsu H., Bartholomew D.P., Lin C.-H. (2005) Foliar application of aviglycine reduces natural flowering in pineapple.
HortScience 40:123-126.
Lin C.-H., Kuan C.-S., Hsu Y.-M., Lin M.-L., Hsu H.-T., Yu C.-W., Bartholomew D.P. (2006) Delaying Natural Flowering in Pineapple. Acta
Horticulturae 702:63-70.
Malezieux E., Zhang J., Sinclair E., Bartholomew D.P. (1994) Predicting pineapple harvest date in different environments, using a computer
simulation model. Agronomy Journal 86:609-617.
Rohrbach K.G., Johnson M. (2003) Pests, diseases and weeds. Pp. 203-251. In: D. P. Bartholomew, et al. (Eds.), The Pineapple: Botany,
Production and Uses, CABI Publishing, Wallingford.
Wang R.H., Maruthasalam S., Lin C.H., Hsu Y.M., Bartholomew D.P. (2007) Delaying Natural Flowering in Pineapple Through Foliar
Application of Aviglycine, an Inhibitor of Ethylene Biosynthesis. HortScience 42:1188-1191.—
38
Newsletter of the Pineapple Working Group, International Society for Horticultural Science
Services
The listings under Commercial Services and Directory of Professionals is maintained as a convenience to readers and should
in no way be construed as an endorsement of those providing commercial or professional services. Those offering specialized
services to pineapple growers or researchers are invited to contact the editor for possible inclusion in the listings below.
Commercial Services
M aintain CF 125 continues to be available for use in pineapple plant propagation. A renewal letter for registration of the product
was received in 2003. For further information, contact Bhushan Mandava, Repar Corporation, P.O. Box 4321, Silver Spring,
MD 20914 Tel: 202-223-1424 Fax: 202-223-0141; E-Mail: mandava@compuserve.com
Centro de Bioplantas. Dr. Justo L. Gonzalez Olmedo, Director of Foreign Affairs Office, Centro De Bioplantas. Universidad De
Ciego De Avila, Carretera a Moron Km 9. Cp69450. Cuba. Centro De Bioplantas offers certificates of authenticity for
pineapple material propagated in their tissue culture facility. W eb site: http://www.Bioplantas.cu
LAM ERSA, Dole's meristem laboratory in Honduras. Contact John T. Mirenda PhD, Dole Fresh Fruit International Ltd., San
Jose, Costa Rica. Phone: 506 287 2175. Fax: 506 287 2675. E-mail: Jmirenda@la.dole.com. The laboratory can produce
meristematically-derived plants of pineapple as well as banana and other crops.
Thai Orchids Lab, Dr. Paiboolya Gavinlertvatana. Horticulture/ agriculture/ forestry tissue culture laboratory with exports to
Australia, U.S.A., Africa, and Asia. MD2 pineapple available (open to acquiring additional varieties) or confidential
exclusive contract propagation. Phone: +1 510 931 7865 Fax: +66 2510 9452 W ebsite: http://www.tolusa.com/ E-mail:
info@tolusa.com.
Vitropic, Zone d'Activités Economiques des Avants, 34270 Saint Mathieu de Tréviers France; Tel: + 33 (0)4 67 55 34 58; Fax: +
33 (0)4 67 55 23 05. E-mail : vitropic@vitropic.fr. W eb site: www.vitropic.fr. Vitropic proposes the best individuals from
the CIRAD FHLOR selected clones including: Cayenne Group, Queen Group, Perolera Group, MD2, Ornamentals
pineapples. The range is continuously extending, do not hesitate to ask for more information.
Professional Services
Mr. Wilbert Campos Alvarado. M.Sc. Tropical Soils & Crop Mgmt. E-mail. wcamposa@gmail.com. Phone: (506) 8815-7271. Apdo. Postal
536-7210, Guapiles, Costa Rica. Experience in all stages of production (soil preparation, plant nutrition, diseases & pest control, PGR use, etc)
of pineapple for the fresh fruit production market as well as experince in packing plant management and in postharvest treatment. Also worked
in pineapple R&D for several years under different climate conditions (Costa Rico, Guatemala, Ecuador).
Ing. Alejandro Chavarría. APDO 4437-56 Pital, San Carlos. Alajuela, Costa Rica. Tel: (506) 88-20-79-55 / (506) 24-73-40-00,
alechava@hotmail.com . I have worked like an International Pineapple Consulting in México, Costa Rica and Brazil. Experienced in project
feasibility, plantation design, agricultural machinery, all aspects of farm crop management, post harvest management and establishment of good
agricultural practices.
Dr. Mark Paul Culik. INCAPER, Rua Alfonso Sarlo 160, CEP 29052-010, Vitoria, ES, Brazil; Tel: 27-3137-9874; markculik3@yahoo.com.
Experience: PhD in Entomology with more than 25 years of agricultural pest management experience in crops ranging from apples to papaya
and pineapple, identification of pests and beneficial arthropods ranging from Collembola to fruit flies, and current work on scale insects with
emphasis on pineapple mealybugs. Areas of specialization: Entomology, Insect and Pest Identification, Integrated Pest Management.
Dr. Francisco Gomez (E-mail: fgomez1@cablecolor.hn) and Jose R. Vasquez, MBA (E-mail: jrva46@excite.com). Golden Pacific Ag
Services, PO.Box 15088, Lomas Miraflores, 4a. Calle, 1a Avenida # 4326, Tegucigalpa, Honduras. Phone: 504 230 1120; 504 969 5568.
Experience: Pineapple and melon production, from seed propagation-planting-field maintenance-forcing-harvesting-post-harvest management
and commercialization.
Mr. Ian Greig. Greig and Associates, P.O. Box 273508, Tampa, FL 33688. Phone: (813) 908-7698; Fax: (813) 963-6229. E-mail:
iang@ag-consult.com. Web site: www.ag-consult.com. Services for all phases of pineapple production but emphasis is on pineapple industry
and market analysis.
Mr. L. Douglas MacClure. 360 Hoopalua Dr., Pukalani, Hawaii, U.S.A. E-mail: norfolkldm@aol.com.
Experience: More than 39 years with Maui Pineapple Company heading plantation and diversified agriculture operations and started the Royal
Coast Tropical Fruit Company in Costa Rica. Collected and summarized production information in Asia and Central America. Also consulted
on pineapple for companies and growers in El Salvador, Australia, Thailand and Indonesia.
39
Newsletter of the Pineapple Working Group, International Society for Horticultural Science
Mr. Graham J. Petty 13 Somerset Place, Lambert Road, Port Alfred, 6170, Republic of South Africa. Phone: +27 (0) 46 624 4868;
Tel/Fax: +27 (0) 46 625 0946; E-mail: grahamp@imaginet.co.za. Experience: M.Sc. (Agric) Pretoria : Pr. Sci. Nat. . Researcher and advisor
to the South African Canning Pineapple Industry on matters of Pest Management in pineapple culture, for 34 years. Economic entomology and
management of biological control agents have received particular attention.
Mr. Col Scott. E-mail: scottch45@bigpond.com. Mobile: +61 488092442; Phone: +61 7 34252417; Fax: +61 7 34252417. Over 37 years
experience in all aspects of pineapple agronomy and research in Australia (32 years with Golden Circle Ltd ) and South Africa (5 years with
Summerpride Foods Ltd). Experience includes working with growers, researchers and fertilizer and agricultural chemical suppliers. Other
production areas visited include Hawaii, Central America, Thailand, Indonesia and Malaysia.
Dr. José Aires Ventura. Incaper, Rua Afonso Sarlo 160 (bento Ferreira), 29052-010, Vitoria-ES, Brazil. E-mail: ventura@incaper.es.gov.br;
Tel.: 55-27-31379874. www.incaper.es.gov.br. Area of Specialization: Plant Pathology (research in pineapple diseases management; Fusarium
diagnosis, diseases resistance).
Mr. Dean Wheeler. AgResults Inc., 609 Buchanan Street, Davis, California, U.S.A. 95616. Phone/fax: 530-758-4620 Residence:
530-758-3354. Email: agresults@aol.com. Web page at http://agresults.com/.
Book Reviews and Web Sites
Book Reviews
No reviews were provided for this issue.
Web Sites of Possible Interest
1.
The CIRAD Market News Service website is http://passionfruit.cirad.fr/index.php/(html)/fruitrop/fruitrop.html. The services
journal FruiTrop can be found at the web site and many publications are available either for a fee and older issues are
available as pdf files at no cost.
2. Abacaxi on-line at http://www.cnpmf.embrapa.br/informativos/abacaxi/abacaxi_online_v5_3_07.pdf
3. Mandioca e Fruticultura Tropical at http://www.cnpmf.embrapa.br/. Publications (in Porutgese) on abacaxi (pineapple) are
available from the Publiçöes link.
4. http://www.redorbit.com/news/science/195814/ntc_marketing_looks_to_expand_appeal_of_its_libbys_brand/
5. http://www.gov.ph/index.php?option=com_content&task=view&id=1962&Itemid=2
6. http://www.freshplaza.com/news_detail.asp?id=24957
7. http://www.laborrights.org/creating-a-sweatfree-world/resources/1758
8. http://www.ars.usda.gov/research/projects/projects.htm?ACCN_NO=410039&showpars=true&fy=2006
9. http://www.ars.usda.gov/research/projects/projects.htm?ACCN_NO=410039&showpars=true&fy=2007
10. http://www.ars.usda.gov/research/projects/projects.htm?ACCN_NO=410039&fy=2008
11. http://www.horticulture.com.au/industry/annualreports.asp
—
References
The list below includes papers related to various aspects of pineapple culture, physiology, processing, preservation or
byproducts that were published or located since the last issue of the newsletter was printed. Some papers may seem relatively
unrelated to pineapple but since judgement must be exercised when including or excluding references, the decision was made to
ere on the side of inclusion so as to serve as many readers as possible. Often, abstracts of the papers listed below can be found online and of course all abstracts of paper published in Acta Horticulturae are available from info@ishs.org.
Pineapple Reference Database
A pineapple references database with over 7,000 references in it is maintained by the editor. Literature searches of the
database on specific topics, including abstracts where available, can be obtained by contacting Duane Bartholomew at
duaneb@hawaii.edu.
Acosta, O., VÃ-quez, F., and Cubero, E., 2008. Optimisation of low calorie mixed fruit jelly by response surface methodology.
Food Quality and Preference 19:79-85.
Aguilar Uscanga, M.G., Escudero Abarca, B.I., Gomez Rodriguez, J., and Cortes Garcia, R., 2007. Carbon sources and their
effect on growth, acetic acid and ethanol production by Brettanomyces bruxellensis in batch culture. Journal of Food Process
Engineering 30:13-23.
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Newsletter of the Pineapple Working Group, International Society for Horticultural Science
Ahmed, O.H., Ahmad, H.M.H., Musa, H.M., Rahim, A.A., and Rastan, S.O.S., 2005. Applied K fertilizer use efficiency in
pineapples grown on a tropical peat soil under residues removal. TheScientificWorldJOURNAL 5:42-49.
Ahmed, O.H., Husni, M.H.A., Hanafi, M.M., Anuar, A.R., and Omar, S.R.S., 2007. Phosphorus fertilizer use in pineapple cultivation
with in situ residues burning on organic soils. Communications in Soil Science and Plant Analysis 38:1243-1254.
Alavi, G., Sanda, M., Loo, B., Green, R.E., and Ray, C., 2008. Movement of bromacil in a Hawaii soil under pineapple cultivation a field study. Chemosphere 72:45-52.
Alejos C., D.R., Mogollón M., N.J., Arizaleta, M., and De Pérez, M., 2005. Evaluación de la Aclimatización y Comportamiento
Hortícola de Plantas de Piña (Ananas comosus L. Merril) 'Queen Australia' Producidas in vitro. Proceedings of the ISTH
49:205-211.
Alves, G.A.R., Lobato, A.K.S., Santos Filho, B.G., Oliveira Neto, C.F., Da Costa, R.C.L., Maia, W.J.M.S., Freitas, J.M.N., and
Silva, L.I., 2008. Interaction among Organic Matter and Pathogen Fusarium subglutinans F. sp. in Soil Cultivated with Ananas
comosus. Agriculture Journal 3:459-462.
Alves, G.A.R. and Nunes, M.A.L., 2008. Survival of Fusarium subglutinans f. sp. ananas in soils. / Sobrevivencia de Fusarium
subglutinans f. sp. ananas em solos. Revista de Ciencias Agrarias:157-171.
Angeles, D.E., 2008. Uptake of P32 in papaya, banana & pineapple grown in a polyculture system. Philippine Journal of Crop
Science 33:69-79.
Anonymous, 2007. Plant variety descriptions submitted for registration of plant breeders' rights in Australia up to 6 March 2008
(Pineapple cultivars Aus-Jubilee and Aus-Carnival; Vol. 20, No. 4 available as a pdf file). Plant Varieties Journal 20:371-371.
Antony, E., Taybi, T., Courbot, M., Mugford, S.T., Smith, J.A.C., and Borland, A.M., 2008. Cloning, localization and expression
analysis of vacuolar sugar transporters in the CAM plant Ananas comosus (pineapple). Journal of Experimental Botany
59:1895-1908.
Antunes, A.M., Ono, E.O., and Sampaio, A.C., 2008. Effect of Paclobutrazol in the control of the natural flowering difference of
'Smooth Cayenne' pineapple plant. / Efeito do Paclobutrazol no controle da diferenciao floral natural do abacaxizeiro cv.
Smooth Cayenne. Revista Brasileira de Fruticultura 30:290-295.
Antunes, A.M., Ono, E.O., Sampaio, A.C., and Rodrigues, J.D., 2008. Physico-chemical and harvest time alterations in pineapple
fruits 'Smooth Cayenne' caused by paclobutrazol. Brazilian Archives of Biology and Technology 51:19-26.
Arabshahi-Delouee, S. and Asna, U., 2007. Application of phenolic extracts from selected plants in fruit juice. International Journal
of Food Properties 10:479-488.
Araujo, R.F., Siqueira, D.L., and Cecon, P.R., 2008. In vitro pineapple shoot proliferation in benzylaminopurine (BAP) and
naphthaleneacetic acid (NAA) concentrations. / Multiplicaio in vitro do abacaxizeiro 'smooth cayenne' utilizando
benzilaminopurina (BAP) e acido naftalenoacetico (ANA). Revista Ceres 55:455-460.
Ashwini, T., Mahesh, B., Jyoti, K., and Uday, A., 2008. Preparation of ferulic acid from agricultural wastes: its improved extraction
and purification. Journal of Agricultural and Food Chemistry 56:7644-7648.
Assawarachan, R. and Noomhorm, A., 2008. Effect of operating condition on the kinetic of color change of concentrated pineapple
juice by microwave vacuum evaporation. Journal of Food, Agriculture & Environment 6:47-53.
Azevedo, B.M.d., Bomfim, G.V.d., Carvalho, A.C.P.P.d., Gondim, R.S., and Viana, T.V.d.A., 2008. Ex vitro acclimatization of
ornamental pineapple under different irrigation levels. / Aclimatizacao ex vitro de abacaxizeiro ornamental com diferentes
laminas de irrigacao. IRRIGA 13:298-309.
Balasundram, S.K., Husni, M.H.A., and Ahmed, O.H., 2008. Application of geostatistical tools to quantify spatial variability of
selected soil chemical properties from a cultivated tropical peat. Journal of Agronomy 7:82-87.
Barguil, B.M., Leocardio Soares Pessoa, W.R., Alves de Oliveira, S.M., and Barbosa Coelho, R.S., 2008. Occurrence of
Pestalotiopsis neglecta in Ananas lucidus. Summa Phytopathologica 34:96.
Bebe, H. and Singh, N.I., 2007. Post-harvest fruit rot of pineapple var. Queen and changes in its ascorbic acid content. Journal of
Mycopathological Research 45:241-244.
Bennici, A. and Bussi, B., 2009. Protocol for an efficient pineapple plant regeneration by in vitro organogenesis and histological
analysis of the morphogenic process. Advances in Horticultural Science 23:49-56.
Bhattacharya, R. and Bhattacharyya, D., 2009. Preservation of natural stability of fruit "bromelain" from Ananas comosus
(pineapple). Journal of Food Biochemistry 33:1-19.
Bhattacharyya, B.K., 2008. Bromelain: an overview. Natural Product Radiance 7:359-363.
Bomfim, G.V.d., Carvalho, A.C.P.P.d., Bezerra, F.C., Azevedo, B.M.d., Viana, T.V.d.A., and Oliveira, K.M.A.S.d., 2007.
Acclimatization of ornamental pineapple micropropagated plants in different substrate volumes. / Aclimatizaco de mudas
micropropagadas de abacaxizeiro ornamental em diferentes volumes de substrato. Revista Brasileira de Horticultura
Ornamental 13:121-128.
Botella, J.R. and Smith, M., 2008. Genomics of pineapple, crowning the king of tropical fruits. In: Moore, P. H. and Ming, R. (eds.),
Genomics of tropical crop plants. Springer-Verlag GmbH, Heidelberg; Germany.
Bregonci, I.d.S., Reis, E.F.d., Almeida, G.D.d., Brum, V.J., and Zucoloto, M., 2008. Evaluation of the foliar and radicular growth of
the micropropagated plantlets of the pineapple cv. Gold in acclimatation. / Avaliação do crescimento foliar e radicular de
mudas micropropagadas do abacaxizeiro cv. Gold em aclimatação. IDESIA Facultad de Agronomía, Universidad de
Tarapacá 26:87-96.
Bregonci, I.d.S., Reis, E.F.d., Almeida, G.D.d., Coelho, R.I., and Brum, V.J., 2008. Foliar contents of macro and micronutrients of
micropropagated seedlings of pineapple Gold in the acclimatization phase with different NPK levels. / Teor foliar de macro e
micronutrientes de mudas micropropagadas de abacaxi Gold na fase de aclimatacao com diferentes niveis de NPK. Revista
Ciencia Agronomica 39:233-239.
Bregonci, I.d.S., Schmildt, E.R., Coelho, R.I., Reis, E.F.d., Brum, V.J., and Santos, J.G.d., 2008. Foliar fertilization with macro and
micronutrients in the growth of plantlets micropropagated of pineapple cv. Gold [Ananas comosus (L.) Merrill] in different
41
Newsletter of the Pineapple Working Group, International Society for Horticultural Science
containers. / Adubacao foliar com macro e micronutrientes no crescimento de mudas micropropagadas do abacaxizeiro cv.
Gold [Ananas comosus (L.) Merrill] em diferentes recipientes. Ciencia e Agrotecnologia 32:705-711.
Brenes-Prendas, S. and Agüero-Alvarado, R., 2007. Weed surveys and identification, and description of their control strategies in
four pineapple (Ananas comosus L.) farms in Costa Rica. / Reconocimiento taxonómico de arvenses y descripción de su
manejo, en cuatro fincas productoras de piña (Ananas comosus L.) en Costa Rica. Agronomia Mesoamericana 18:239-246.
Brito, C.A.K.d., Sato, H.H., Spironello, A., and Siqueira, W.J., 2007. IAC Gomo-de-mel pineapple (Ananas comosus (L.) Merrill):
characteristics of pulp and peroxidase of juice. / Abacaxi IAC Gomo-de-mel (Ananas comosus (L.) Merrill): Características da
polpa e da peroxidase do suco. Boletim do Centro de Pesquisa e Processamento de Alimentos 25:257-266.
Brito Neto, J.F.d., Sá Sobrinho, R.G.d., Pereira, W.E.d., Barbosa, J.A., S. Costa, D.d., Lacerda, J.S., dos Santos, D.P., and O.
Vieira, D.d., 2009. Commercialization forms and organization of pineapple producers in the state of Paraíba, Brazil
Acta Horticulturae 322:313-316.
Buzrul, S., Alpas, H., Largeteau, A., and Demazeau, G., 2008. Inactivation of Escherichia coli and Listeria innocua in kiwifruit and
pineapple juices by high hydrostatic pressure. International Journal of Food Microbiology 124:275-278.
Carvalho, L.M.J.d., Castro, I.M.d., and Silva, C.A.B.d., 2008. A study of retention of sugars in the process of clarification of
pineapple juice (Ananas comosus, L. Merril) by micro- and ultra-filtration. Journal of Food Engineering 87:447-454.
Catunda, P.H.A., Marinho, C.S., Assis Gomes, M.M.d., and Cordeiro de Carvalho, A.J., 2008. Brassinosteroid and substrates in
acclimatization of 'Imperial' pineapple. Acta Scientiarum Agronomy 30:345-352.
Chaisakdanugull, C., Theerakulkait, C., and Wrolstad, R.E., 2007. Pineapple juice and its fractions in enzymatic browning
inhibition of banana [Musa (AAA group) Gros Michel]. Journal of Agricultural and Food Chemistry 55:4252-4257.
Charanjiv, S., Sharma, H.K., and Sarkar, B.C., 2008. Optimization of process conditions during osmotic dehydration of fresh
pineapple. Journal of Food Science and Technology (Mysore) 45:312-316.
Chen, H., Huang, T., Tsai, C., and Mujumdar, A.S., 2008. Development and performance analysis of a new solar energy-assisted
photocatalytic dryer. Drying Technology 26:503-507.
Cheng, K., Wu, Q., Zheng, Z., Peng, X., Simon, J.E., Chen, F., and Wang, M., 2007. Inhibitory effect of fruit extracts on the
formation of heterocyclic amines. Journal of Agricultural and Food Chemistry 55:10359-10365.
Chien, S. and Kang, P., 2008. Functional and baking properties of pineapple core for bread making. Taiwanese Journal of
Agricultural Chemistry and Food Science 46:7-13.
Chonhenchob, V., Kamhangwong, D., and Singh, S.P., 2008. Comparison of reusable and single-use plastic and paper shipping
containers for distribution of fresh pineapples. Packaging Technology & Science 21:73-83.
Chonratamarit, S., Thaipakdee, S., Chutvachiravong, P., and Damrongkittikul, S., 2008. Development of web-based information
systems of economic field crops in the southern Thailand. Kasetsart Journal, Social Sciences 29:67-75.
Coelho, R.I., Carvalho Júnior, A.C.d., Lopes, J.C., Teixeira, S.L., and Marinho, C.S., 2007. 'Smooth Cayenne' pineapple crown on
the yield of suckers type planting material. / Coroa do abacaxi 'Smooth Cayenne' na produção de mudas do tipo rebentão.
Ciencia e Agrotecnologia 31:1867-1871.
Coelho, R.I., Lopes, J.C., Cordeiro de Carvalho Júnior, A., Amaral, J.A.T.d., and Matta, F.d.P., 2007. Nutritional status and
growth caracteristics of pineapple in dystrophic yellow latosol cultivated in function of NPK fertilization. / Estado nutricional e
características de crescimento do abacaxizeiro 'Jupi' cultivado em latossolo amarelo distrófico em função da adubação com
NPK. Ciencia e Agrotecnologia 31:1696-1701.
Conley, T.G. and Udry, C.R., 2000. Learning about a New Technology: Pineapple in Ghana. Economic Growth Center, Yale
University, P.O. Box 208269, New Haven, CT;http://www.econ.yale.edu/~egcenter/
http://www.econ.yale.edu/growth_pdf/cdp817.pdf, New Haven.
Conley, T.G. and Udry, C.R., 2005. Learning about a New Technology: Pineapple in Ghana. 63 pages. Economic Growth Center,
Yale University, P.O. Box 208269, New Haven, CT;http://www.econ.yale.edu/~cru2//pdf/july2005a.pdf, New Haven.
Costa, D.M.d., Zahra, A.R.F., Kalpage, M.D., and Rajapakse, E.M.G., 2008. Effectiveness and molecular characterization of
Burkholderia spinosa, a prospective biocontrol agent for controlling postharvest diseases of banana. Biological Control
47:257-267.
da Conceicao Pedrera, A.C., Naves, R.V., and do Nascimento, J.L., 2008. Seasonal variation of the pineapple cv Perola quality in
Goiania, Goias State, Brazil. Pesquisa Agropecuaria Tropical 38:262-268.
Dacera, D.d.M. and Babel, S., 2008. Removal of heavy metals from contaminated sewage sludge using Aspergillus niger
fermented raw liquid from pineapple wastes. Bioresource Technology 99:1682-1689.
Danso, K.E., Ayeh, K.O., Oduro, V., Amiteye, S., and Amoatey, H.M., 2008. Effect of 6-benzylaminopurine and α-naphthalene
acetic acid on in vitro production of MD2 pineapple planting materials. World Applied Sciences Journal 3:614-619.
Davey, M.R., Sripaoraya, S., Anthony, P., Lowe, K.C., and Power, J.B., 2007. Pineapple. In: Pua, E. C. and Davey, M. R. (eds.),
Transgenic Crops V. Springer-Verlag GmbH, Heidelberg; Germany.
de Carvalho Correia, M.X., Costa, R.G., da Silva, J.H.V., de Carvalho, F.F.R., and de Medeiros, A.N., 2006. Use of dehydrated
pineapple by-product in diets for growing goats: digestibility and performance. Revista Brasileira de Zootecnia 35:1822-1828.
de Carvalho, L.M.J., da Silva, C.A.B., and de Castro, I.M., 2008. A study of retention of sugars in the process of clarification of
pineapple juice (Ananas comosus, L. Merril) by micro- and ultra-filtration [electronic resource]. Journal of Food Engineering
87:447-454.
De Silva, A.E., Kadir, M.A., Aziz, M.A., and Kadzimin, S., 2006. High and rapid callus proliferation of pineapple (Ananas comosus
L.) cv. Moris. TheScientificWorldJOURNAL 6:169-175.
Deka, B.C., Sharma, S., Choudhury, S., and Pal, R.K., 2008. Development of a packaging system for distant transportation of
'Kew' pineapple (Ananas comosus). Indian Journal of Agricultural Sciences 78:340-342.
Dhar, M., Rahman, S.M., and Sayem, S.M., 2008. Maturity and post harvest study of pineapple with quality and shelf life under red
soil. International Journal of Sustainable Crop Production 3:69-75.
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Newsletter of the Pineapple Working Group, International Society for Horticultural Science
Eduardo, M.d.P., Benedetti, B.C., and Ferraz, A.C.O., 2008. Firmness indexes evaluation for fresh-cut sliced pineapple treated
with calcium salts solutions. / Avaliação de Ã-ndices de firmeza para abacaxi minimamente processado em fatias tratadas
com soluções de sais de cálcio. Engenharia AgrÃ-cola 28:154-163.
Eduardo Ordonez-Santos, L., Isabel Gonzalez-Loaiza, D., and Andres Ospina-Aragon, C., 2008. Comment to the paper by
Chutintrasri and Noomhorm (2006), Thermal inactivation of polyphenoloxidase in pineapple puree, LWT 39 (2006) 492-495.
LWT - Food Science and Technology 41:1739.
Elias Junior, J., Gomes, D.C., Matos, A.P.d., and Almeida, C.O.d., 2009. Micro and macroeconomic analyses of the pineapple
industry in the state of Tocantins
Acta Horticulturae 322:317-322.
Fan, X. and Sokorai, K.J.B., 2008. Effect of ionizing radiation on furan formation in fresh-cut fruits and vegetables. Journal of Food
Science 73:C79-C83.
Faure, G., Veerabadren, S., and Hocdé, H., 2008. Family agriculture under standards: a challenge for the pineapple farmers in
Costa Rica? / L'agriculture familiale mise sous normes: un défi pour les producteurs d'ananas au Costa Rica? Économie
Rurale:184-197.
Fernandes, F.A.N., Gallão, M.I., and Rodrigues, S., 2008. Effect of osmosis and ultrasound on pineapple cell tissue structure
during dehydration. Journal of Food Engineering 90:186-190.
Fernandes, F.A.N. and Rodrigues, S., 2008. Application of ultrasound and ultrasound-assisted osmotic dehydration in drying of
fruits. Drying Technology 26:1509-1516.
Ferreira, A.C.H., Rodriguez, N.M., Neiva, J.N.M., Campos, W.E., and Borges, I., 2007. Bromatologic and fermentative
characteristics of silages of napier grass mixed with increasing levels of pineapple industrial residues. / CaracterÃ-sticas
quimico-bromatológicas e fermentativas do capim-elefante ensilado com nÃ-veis crescentes de subproduto da
agroindústria do abacaxi. Revista Ceres 54:99-107.
Gambley, C.F., Geering, A.D.W., Steele, V., and Thomas, J.E., 2008. Identification of viral and non-viral reverse transcribing
elements in pineapple (Ananas comosus), including members of two new badnavirus species. Archives of virology
153:1599-1604.
Gambley, C.F., Geering, A.D.W., and Thomas, J.E., 2009. Development of an immunomagnetic capture-reverse
transcriptase-PCR assay for three pineapple ampeloviruses. Journal of Virological Methods 155:187-192.
Gambley, C.F., Steele, V., Geering, A.D.W., and Thomas, J.E., 2008. The genetic diversity of ampeloviruses in Australian
pineapples and their association with mealybug wilt disease. Australasian Plant Pathology 37:95-105.
Ghosh, R., Chakraborty, J., and Ghosh, D., 2008. Peroxidase activity of two cultivars (Kew and Queen) of ripe pineapple (Ananas
comosus) of Tripura. Journal of Applied Bioscience 34:106-109.
Glenn, D.M., Wuensche, J., McIvor, I., Nissen, R., and George, A., 2008. Ultraviolet radiation effects on fruit surface respiration
and chlorophyll fluorescence. Journal of Horticultural Science & Biotechnology 83:43-50.
Gnanasoundari, S., Sashi, V., Malathy, N.S., and Vasantha, S., 2007. Xylanase production from Aspergillus flavus. Plant Archives
7:213-214.
Gnanavelrajah, N., Shrestha, R.P., Schmidt-Vogt, D., and Samarakoon, L., 2008. Carbon stock assessment and soil carbon
management in agricultural land-uses in Thailand. Land Degradation & Development 19:242-256.
Gupta, P., Maqbool, T., and Saleemuddin, M., 2007. Oriented immobilization of stem bromelain via the lone histidine on a metal
affinity support. Journal of Molecular Catalysis B Enzymatic 45:78-83.
Habib, S., Khan, M.A., and Younus, H., 2007. Thermal destabilization of stem bromelain by trehalose. Protein Journal 26:117-124.
Hale, L.P., Fitzhugh, D.J., and Staats, H.F., 2006. Oral immunogenicity of the plant proteinase bromelain. International
Immunopharmacology 6:2038-2046.
Hamad, A.M. and Rosna Mat, T., 2008. The effect of different hormones and incubation periods on in vitro proliferation of
pineapple (Ananas comosus L.) Merr cv. Smooth cayenne shoot-tip culture. Pakistan Journal of Biological Sciences
11:386-391.
Hamad, A.M. and Rosna Mat, T., 2008. Effect of sequential subcultures on in vitro proliferation capacity and shoot formations
pattern of pineapple (Ananas comosus L. Merr.) over different incubation periods. Scientia Horticulturae 117:329-334.
Hamad, A.M. and Taha, R.M., 2008. Effect of benzylaminopurine (BAP) on in vitro proliferation and growth of pineapple (Ananas
comosus L. Merr.) cv. Smooth Cayenne. Journal of Applied Sciences 8:4180-4185.
Hamed, A.M. and Ali, E.A.M., 2007. Effect of different sea water concentrations on growth parameters of pineapple (Ananas
comosus) in vitro and in vivo. Journal of Applied Sciences Research:713-722.
Hameed, B.H., Krishni, R.R., and Sata, S.A., 2009. A novel agricultural waste adsorbent for the removal of cationic dye from
aqueous solutions. Journal of Hazardous Materials 162:305-311.
Hasan, S.M.Z. and Abdullah, N.S., 2007. Effect of salinity on growth, proline accumulation and malate content of pineapple
(Ananas comosus (L.) Merill.) under tissue culture condition. Malaysian Applied Biology 36:57-63.
He, Y., Luo, J., Hu, Z., Wang, R., Gao, A., Fang, S., Zhao, C., Yang, X., Ye, Z., and Wang, Z., 2008. Classification and
organogenesis of non-embryogenic callus from Ananas comosus. Journal of Fruit Science 25:65-68.
He, Y., Luo, J., Wu, H., Wang, R., Gao, A., Zhao, C., Yu, X., Ye, Z., Wang, Z., Han, J., and Liu, H., 2007. Somatic embryogenesis
from leaf base callus of Ananas comosus. Journal of Fruit Science 24:59-63.
Hebbar, H.U., Sumana, B., and Raghavarao, K.S.M.S., 2008. Use of reverse micellar systems for the extraction and purification of
bromelain from pineapple wastes. Bioresource Technology 99:4896-4902.
Henson, I.E., 2007. Plantations on peat: how sustainable are they? Environmental aspects of developing peat lands for
agriculture. Planter 83:21-39.
Hernandez, Y., Lobo, M.G., and Gonzalez, M., 2009. Factors affecting sample extraction in the liquid chromatographic
determination of organic acids in papaya and pineapple. Food Chemistry 114:734-741.
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Newsletter of the Pineapple Working Group, International Society for Horticultural Science
Hidaka, M., Nagata, M., Kawano, Y., Sekiya, H., Kai, H., Yamasaki, K., Okumura, M., and Arimori, K., 2008. Inhibitory effects of
fruit juices on cytochrome P450 2C9 activity in vitro. Bioscience, Biotechnology and Biochemistry 72:406-411.
Hongvaleerat, C., Cabral, L.M.C., Dornier, M., Reynes, M., and Ningsanond, S., 2008. Concentration of pineapple juice by osmotic
evaporation. Journal of Food Engineering 88:548-552.
Hou, C., Lin, Y., Wang, Y., Jiang, C., and Wu, M., 2008. Effect of storage conditions on methanol content of fruit and vegetable
juices. Journal of Food Composition and Analysis 21:410-415.
Hseu, S.H., Zeng, W.F., Lai, W.C., Pan, Y.P., and Lin, C.Y., 2008. Fruit rot disease of pineapple caused by Burkholderia gladioli.
Plant Pathology Bulletin 17:157-167.
Imandi, S.B., Bandaru, V.V.R., Somalanka, S.R., Bandaru, S.R., and Garapati, H.R., 2008. Application of statistical experimental
designs for the optimization of medium constituents for the production of citric acid from pineapple waste. Bioresource
Technology 99:4445-4450.
Inclan, D.J., Bermudez, F.J., Alvarado, E., Ellis, M., Williams, R.N., and Acosta, N., 2008. Comparison of biological and
conventional insecticide treatments for the management of the pineapple fruit borer, Strymon megarus (Lepidoptera:
Lycaenidae) in Costa Rica. Ecological Engineering 34:328-331.
Jacques, D.T., Marc, K.T., and Marius, A., 2008. Biochemical effectiveness in liver detoxication of fresh pineapple (Ananas
comosus) with the wistar rats, previously intoxicated by Doliprane®. Journal of Cell and Animal Biology 2:31-35.
Jaeger de Carvalho, L.M., de Castro, I.M., and Bento da Silva, C.A., 2008. A study of retention of sugars in the process of
clarification of pineapple juice (Ananas comosus, L. Merril) by micro- and ultra-filtration. Journal of Food Engineering
87:447-454.
Jai, P., Singh, N.P., and Sankaran, M., 2008. Influence of plant population on fruiting, fruit quality and yield of pineapple (Annanas
comosus) under tropical tilla land of Tripura. Indian Journal of Agricultural Sciences 78:801-802.
Jamal, P., Alam, M.Z., and Suhani, F., 2008. Biophenols from agro-industrial wastes. Medical Journal of Malaysia 63:107-108.
Jetana, T., Kitsamraj, S., Sophon, S., Tasripoo, K., and Vongpipatana, C., 2009. The comparative study digestion and metabolism
of nitrogen and purine derivatives in male, Thai, Swamp buffalo and Thai, Brahman cattle [electronic resource]. Animal
Science Journal 80:130-139.
Jetana, T., Sophon, S., Liang, J.B., Vongpipatana, C., Suthikrai, W., and Usawang, S., 2009. The effects of concentrate added to
pineapple (Ananas Comosus linn. Mer.) waste silage in differing ratios to form complete diets, on digestion, excretion of
urinary purine derivatives and blood metabolites in growing, male, Thai swamp buffaloes [electronic resource]. Tropical
animal health and production 41:449-459.
Jirasripongpun, K., Pewlong, W., Kitraksa, P., and Krudngern, C., 2008. Carotenoid production by Xanthophyllomyces
dendrorhous: use of pineapple juice as a production medium. Letters in Applied Microbiology 47:112-116.
Jose, J.S., Montes, R., and Nikonova, N., 2007. Seasonal patterns of carbon dioxide, water vapour and energy fluxes in
pineapple. Agricultural and Forest Meteorology 147:16-34.
Joseph, T.S. and Mathew, M., 2007. Economics and cultivation of pineapple in Kerala. Journal of Economic and Taxonomic
Botany 31:990-995.
Joyeeta, S., Ruma, G., and Ghosh, D., 2008. Biochemical constituents of two cultivars of ripe pineapple (Ananas comosus) of
Tripura. Journal of Applied Bioscience 34:91-93.
Jung, Y., Choi, C., Park, J., Kang, H., Choi, J., Nou, I., Lee, S., and Kang, K., 2008. Overexpression of the pineapple fruit
bromelain gene (BAA) in transgenic Chinese cabbage (Brassica rapa) results in enhanced resistance to bacterial soft rot.
EJB, Electronic Journal of Biotechnology 11:Article 7-Article 7.
Jutamongkon, R. and Charoenrein, S. 2008. The relationship between bromelain activity and total soluble solids in Smooth
Cayenne type pineapple, Bangkok; Thailand.
Kaewtathip, T. and Charoenrein, S. 2008. Effect of packaging and freezing rate on off-odor in frozen pineapple, Bangkok;
Thailand.
Kaneshiro, W.S., Burger, M., Vine, B.G., de Silva, A.S., and Alvarez, A.M., 2008. Characterization of Erwinia chrysanthemi from a
bacterial heart rot of pineapple outbreak in Hawaii. Plant Disease 92:1444-1450.
Kapoor, I.P.S., Bandana, S., and Gurdip, S., 2008. Essential oil and oleoresins of Cinnamomum tamala (Tejpat) as natural food
preservatives for pineapple fruit juice. Journal of Food Processing and Preservation 32:719-728.
Keyser, M., Muller, I.A., Cilliers, F.P., Nel, W., and Gouws, P.A., 2008. Ultraviolet radiation as a non-thermal treatment for the
inactivation of microorganisms in fruit juice. Innovative Food Science & Emerging Technologies 9:348-354.
Khatoon, H., Younus, H., and Saleemuddin, M., 2007. Stem bromelain: An enzyme that naturally facilitates oriented
immobilization. Protein and Peptide Letters 14:233-236.
Kole, C., 2007. Fruits and nuts. Genome Mapping and Molecular Breeding in Plants, (one of 7 volumes that provide a timely
overview of the current status of genome analysis in plants including pineapple)
In: Kole, C. (ed.), Fruits and nuts. Springer-Verlag GmbH, Heidelberg; Germany.
Korbin, M., 2008. Genetic modification of fruit plants - directions, successes and obstacles. Biotechnologia (Poznan):9-19.
Koutchma, T., Paris, B., and Patazca, E., 2007. Validation of UV coiled tube reactor for fresh juices. Journal of Environmental
Engineering and Science 6:319-328.
Kurosumi, A., Sasaki, C., Yamashita, Y., and Nakamura, Y., 2009. Utilization of various fruit juices as carbon source for production
of bacterial cellulose by Acetobacter xylinum NBRC 13693. Carbohydrate Polymers 76:333-335.
Lapez-Malo, A. and Palou, E., 2008. Storage stability of pineapple slices preserved by combined methods. International Journal of
Food Science & Technology 43:289-295.
Leong, Y., Xui, O., and Chia, O., 2008. Survival of SA11 rotavirus in fresh fruit juices of pineapple, papaya, and honeydew melon.
Journal of Food Protection 71:1035-1037.
Liao, J., Hu, C., Chang, C., and Deng, T., 2008. The preliminary identification of Pineapple mealybug wilt-associated virus-1 on
pineapple in Taiwan. Journal of Taiwan Agricultural Research 57:1-14.
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Newsletter of the Pineapple Working Group, International Society for Horticultural Science
Lin, Q., Wang, Y.M., Nose, A., Hong, H.T.K., and Agarie, S., 2008. Effects of high night temperature on lipid and protein
compositions in tonoplasts isolated from Ananas comosus and Kalanchoe pinnata leaves. Biologia Plantarum 52:59-65.
Little, H.A., Grumet, R., and Hancock, J.F., 2009. Modified Ethylene Signaling as an Example of Engineering for Complex Traits:
Secondary Effects and Implications for Environmental Risk Assessment. HortScience 44:94-101.
Liu, C., Hsu, C., and Hsu, M., 2007. Improving the quality of fresh-cut pineapples with ascorbic acid/sucrose pretreatment and
modified atmosphere packaging. Packaging Technology & Science 20:337-343.
Liu, W., Yi, G., Liu, Y., Zhang, Q., and Zeng, J., 2008. Identification of pineapple germplasm and relationship analysis with AFLP.
Journal of Fruit Science 25:516-520.
Liu, Y., Yi, Q., Zhong, Y., Pan, J., Meng, X., and Liu, C., 2007. The performance of Australian Kayin pineapple cultivar in
Guangzhou area, Guangdong province. China Fruits:28-30.
Lopez-Malo, A. and Palou, E., 2008. Storage stability of pineapple slices preserved by combined methods. International Journal of
Food Science & Technology 43:289-295.
Lorenzo, J.C. and Garcia-Borroto, M., 2008. Use of regression analysis in plant cell, tissue, and organ culture experiments. In
Vitro Cellular & Developmental Biology - Plant 44:229-232.
Lorsuwan, P., Rechtanapun, C., and Chantanawarangoon, S. 2008. Total phenolics, radical scavenging capacity and antimicrobial
property of fruit peels, Bangkok; Thailand.
Luan, F., Liu, H.T., Wen, Y.Y., and Zhang, X.Y., 2008. Classification of the fragrance properties of chemical compounds based on
support vector machine and linear discriminant analysis. Flavour and Fragrance Journal 23:232-238.
Ma, C., Xiao, S.-Y., Li, Z.-G., Wang, W., and Du, L.-J., 2007. Characterization of active phenolic components in the ethanolic
extract of Ananas comosus L. leaves using high-performance liquid chromatography with diode array detection and tandem
mass spectrometry. Journal of Chromatography A 1165:39-44.
Maheswarappa, H.P., 2008. In-situ waste management in integrated nutrient management system under coconut (Cocos
nucifera)-based high density multi-species cropping system in tropical soils of India. Indian Journal of Agricultural Sciences
78:924-928.
Majumdar, B., Venkatesh, M.S., and Saha, R., 2007. Long-term effect of various agroforestry systems on soil characteristics and
forms of nitrogen build up in acidic alfisol of Meghalaya. Environment and Ecology 25:798-803.
Makkar, H.P.S., Francis, G., and Becker, K., 2007. Bioactivity of phytochemicals in some lesser-known plants and their effects and
potential applications in livestock and aquaculture production systems. Animal 1:1371-1391.
Manish, D., Verma, R.C., and Jain, M.K., 2008. Mathematical models for prediction of rheological parameters of pineapple juice.
International Journal of Food Engineering 4:article 3-article 3.
Mauler Júnior, J., Silva, P.B.L.d., Calçada, L.A., and Scheid, C.M., 2008. The drying process using an aeolian exhaust fan. /
Secagem utilizando exaustor eólico. Revista Brasileira de Armazenamento 33:29-42.
Medda, P.S., Maitra, S., and Singh, L.S., 2008. Favourable effect of intercropping on the growth of young coconut plantation.
Indian Coconut Journal 38:13-15.
Melo, B.d., Galvão, S.R.A.A., Lopes, P.S.N., Silva, A.P.P.d., Martins, M., Santana, J.d.G., and Luz, J.M.Q., 2007. D-leaf length
and ethephon doses on some characteristics of Smooth Cayenne pineapple in Triângulo Mineiro. / Doses de ethephon e
comprimentos de folhas d sobre algumas caracteristicas do abacaxizeiro, cv Smooth Cayenne no Triângulo Mineiro.
Bioscience Journal 23:7-13.
Melo, E.d.A., Maciel, M.I.S., Lima, V.L.A.G.d., and Nascimento, R.J.d., 2008. Antioxidant capacity of the fruits. / Capacidade
antioxidante de frutas. Revista Brasileira de Ciências Farmacêuticas 44:193-201.
Melzer, M.J., Sether, D.M., Karasev, A.V., Borth, W., and Hu, J.S., 2008. Complete nucleotide sequence and genome
organization of pineapple mealybug wilt-associated virus-1. Archives of virology 153:707-714.
Mendiola, J.A., Marin, F.R., Señoráns, F.J., Reglero, G., MartÃ-n, P.J., Cifuentes, A., and Ibáñez, E., 2008. Profiling of
different bioactive compounds in functional drinks by high-performance liquid chromatography. Journal of Chromatography, A
1188:234-241.
Mezadri, T., Bramorski, A., Faria, M.P.d., Cunha, T.I.d., and Costa, A.d.A.d.S.d., 2008. Total polyphenol index and vitamin C
concentration in fruit pulps produced in Santa Catarina, Brazil. / Õ ndice de polifenóis totais e concentração de vitamina
C em polpas de frutas produzidas em Santa Catarina, Brasil. Higiene Alimentar 22:22-25.
Miguel, A.C.A., Spoto, M.H.F., Abrahão, C., and Silva, P.P.M.d., 2007. Consumer profile evaluation by quality function
development for a pineapple. / Aplicação do método QFD na avaliação do perfil do consumidor de abacaxi pérola.
Ciência e Agrotecnologia 31:563-569.
Mitchell, A.M., Strobel, G.A., Hess, W.M., Vargas, P.N., and Ezra, D., 2008. Muscodor crispans, a novel endophyte from Ananas
ananassoides in the Bolivian Amazon. Fungal Diversity 31:37-43.
Mitchell, S.A. and Ahmad, M.H., 2007. Medicinal plant biotechnology research in Jamaica - challenges and opportunities. Acta
Horticulturae:171-181.
Mohamed, A.R., Sapuan, S.M., Shahjahan, M., and Khalina, A., 2009. Characterization of pineapple leaf fibers from selected
Malaysian cultivars. Journal of Food, Agriculture & Environment 7:235-240.
Montero-Calderón, M., Rojas-Graü, M.A., and MartÃ-n-Belloso, O., 2008. Effect of packaging conditions on quality and
shelf-life of fresh-cut pineapple (Ananas comosus). Postharvest Biology and Technology 50:182-189.
Montilla, R., GarcÃ-a, J.L., Lacruz, L., and Duran, D., 2007. Spalangia drosophilae Ashmead (Hymenoptera: Pteromalidae)
parasitic of pupas of the fly of fragmentation Melanoloma viatrix Hendel (DÃ-ptera: Richardiidae) in Trujillo, Venezuela. /
Spalangia drosophilae Ashmead (Hymenoptera: Pteromalidae) parasitoide de pupas de la mosca de la pina Melanoloma
viatrlx Hendel (DÃ-ptera: Richardiidae) en Trujillo, Venezuela. AgronomÃ-a Tropical (Maracay) 57:107-112.
Morales-Payan, J.P., Marquez-Mendez, P.E., and Martinez-Garrastazu, S.L., 2008. Pineapple fruit and propagule yield as
affgected by a seaweed extract, kinetin, and chlorflurenol (abstract). HortScience 43:1293.
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Newsletter of the Pineapple Working Group, International Society for Horticultural Science
Moreira, M.A., Fraguas, C.B., de Carvalho, J.G., and Pasqual, M., 2007. Micropropagation of pineapple cv. Perola with urea as
nitrogen source. Acta Scientiarum Agronomy 29:689-693.
Munawar, S.S., Umemura, K., and Kawai, S., 2007. Characterization of the morphological, physical, and mechanical properties of
seven nonwood plant fiber bundles. Journal of Wood Science 53:108-113.
Naik, S.T. and Maheswarappa, V., 2007. Prospects of using plant extracts in management of pineapple heart rot. Karnataka
Journal of Agricultural Sciences 20:180-182.
Nakamura, T., Yano, Y., Maruyama, A., and Koh, K., 2008. An analysis of wildlife damage to pineapple production: a case of
Higashi Village, Kunigami County. Journal of Rural Economics:280-287.
Nelson, E.A. and Sage, R.F., 2008. Functional constraints of CAM leaf anatomy: tight cell packing is associated with increased
CAM function across a gradient of CAM expression. Journal of Experimental Botany 59:1841-1850.
Nunes, M.C.N., Emond, J.P., Rauth, M., Dea, S., and Chau, K.V., 2009. Environmental conditions encountered during typical
consumer retail display affect fruit and vegetable quality and waste. Postharvest Biology and Technology 51:232-241.
Obi, O.O., Omole, A.J., Ajasin, F.O., and Tewe, O.O., 2008. Nutritive potentials of four conventional forages fed to growing
grass-cutter (Thryonomys swinderianus). Livestock Research for Rural Development 20:179-179.
Obire, O., Ramesh, R.P., Dick, A.A., and Okigbo, R.N., 2008. Biotechnology influence for the production of ethyl alcohol (ethanol)
from waste fruites. e-Journal of Science & Technology 3:17-32.
Ochoa-MartÃnez, C.I., Ramaswamy, H.S., and Ayala-Aponte, A.A., 2007. Artificial neural network modeling of osmotic
dehydration mass transfer kinetics of fruits. Drying Technology 25:85-95.
October, 2008. Pantoea citrea. [Distribution map], p. Ma-Ma, Distribution Maps of Plant Diseases. CABI, Wallingford; UK.
Oliveira, E.C.P.d., Lameira, O.A., Sousa, F.I.B.d., and Silva, R.J.F., 2008. Leaf structure of curaua in different intensities of
photosynthetically active radiation. / Estrutura foliar de curauá em diferentes intensidades de radiação
fotossinteticamente ativa. Pesquisa Agropecuária Brasileira 43:163-169.
Omotoso, S.O. and Akinrinde, E.A., 2007. Influence of propagule weights and nitrogen fertilizer rates on growth and yield of
pineapple (Ananas comosus (L.) Merr). Asian Journal of Agricultural Research 1:131-136.
Omoya, F.O. and Akharaiyi, F.C., 2008. Studies on qualitative and quantitative characterization of alcoholic beverages from
tropical fruits. Research Journal of Microbiology 3:429-435.
Pasqual, M., Santos, F.C., Figueiredo, M.A.d., Junqueira, K.P., Rezende, J.C.d., and Ferreira, E.A., 2008. Protocol for in vitro
micropropagation of ornamental pineapple. / Micropropagação do abacaxizeiro ornamental. Horticultura Brasileira
26:45-49.
Pathaveerat, S., Terdwongworakul, A., and Phaungsombut, A., 2008. Multivariate data analysis for classification of pineapple
maturity. Journal of Food Engineering 89:112-118.
Pe̕ rez, G., Lorenzo, J.C., Isidro̕ n, M., and Yanes, E., 2009. Phenotypic and AFLP characterization of two new pineapple
somaclones derived from in vitro culture [electronic resource]. Plant cell, tissue, and organ culture 96:113-116.
Peiro-Mena, R., Camacho, M.M., and Martinez-Navarrete, N., 2007. Compositional and physicochemical changes associated to
successive osmodehydration cycles of pineapple (Ananas comosus). Journal of Food Engineering 79:842-849.
Pereira, F.D., Brasil Pereira Pinto, J.E., Silva Rosadol, L.D., Melo de Castro, D., de Arruda Rodrigues, H.C., Beijo, L.A., and
Lameira, O.A., 2007. Anatomical characters of curaua foliar fibers from plantlets in vitro propagation. Acta Scientiarum
Biological Sciences 29:23-28.
Pereira, F.D., Pinto, J.E.B.P., Rosado, L.D.S., Rodrigues, H.C.A., Bertolucci, S.K.V., and Lameira, O.A., 2008. Micropropagation
of the fiber-rich Amazonian species Ananas erectifolius (Bromeliaceae). HortScience 43:2134-2137.
Perez, G., Yanes, E., Isidron, M., and Lorenzo, J.C., 2009. Phenotypic and AFLP characterization of two new pineapple
somaclones derived from in vitro culture. Plant Cell Tissue and Organ Culture 96:113-116.
Perez-Tinoco, M.R., Perez, A., Salgado-Cervantes, M., Reynes, M., and Vaillant, F., 2008. Effect of vacuum frying on main
physicochemical and nutritional quality parameters of pineapple chips. Journal of the Science of Food and Agriculture
88:945-953.
Pineiro, M. and Diaz Rios, L.B., 2007. Implementing programmes to improve safety and quality in fruit and vegetable supply
chains: benefits and drawbacks. Latin American case studies. In: Pineiro, M. and Diaz Rios, L. B. (eds.), Implementing
programmes to improve safety and quality in fruit and vegetable supply chains: benefits and drawbacks. Latin American case
studies. Food and Agriculture Organization of the United Nations (FAO), Rome; Italy.
Prabhakar, S., Abha, S., Ranjana, S., and Singh, A.K., 2007. Utilization of guava juice by value addition through blended
beverages. Acta Horticulturae:639-645.
Prakash, J., Singh, N.P., and Sankaran, M., 2008. Influence of plant population on fruiting, fruit quality and yield of pineapple
(Annanas comosus) under tropical tilla land of Tripura. Indian Journal of Agricultural Sciences 78:801-802.
Rabovich, C., Paull, R., and Sipes, B., 2009. Protease inhibitors and reproduction of reniform nematode in pineapple. Annals of
Applied Biology 154:127-132.
Raja, R.D.A., Bai, R.R., and Prakash, J.W., 2008. Analytical estimation of ascorbic acid in some fruits and vegetables in different
cooking methods. Plant Archives 8:203-205.
Ramadan, M.F., Osman, A.O.M., and El-Akad, H.M., 2008. Total antioxidant potential of juices and beverages: screening by
DPPH in vitro assay. Deutsche Lebensmittel-Rundschau 104:235-239.
Ramsaroop, R.E.S. and Saulo, A.A., 2007. Comparative consumer and physicochemical analysis of Del Monte Hawai'i Gold and
Smooth Cayenne pineapple cultivars. Journal of Food Quality 30:135-159.
Ravelo-Perez, L.M., Hernandez-Borges, J., and Angel Rodriguez-Delgado, M., 2008. Multi-walled carbon nanotubes as efficient
solid-phase extraction materials of organophosphorus pesticides from apple, grape, orange and pineapple fruit juices. Journal
of Chromatography A 1211:33-42.
Reinhardt, A. and Rodriguez, L.V., 2009. Industrial processing of pineapple - trends and perspectives. Acta Horticulturae
822:323-328.
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Newsletter of the Pineapple Working Group, International Society for Horticultural Science
Rio, J.C.d., Rencoret, J., Marques, G., Gutierrez, A., Ibarra, D., Santos, J.I., Jiminez-Barbero, J., Zhang, L., and Martinez, Ã.T.,
2008. Highly acylated (acetylated and/or p-coumaroylated) native lignins from diverse herbaceous plants. Journal of
Agricultural and Food Chemistry 56:9525-9534.
Robert, S.D., Aziz Al-safi, I., Than, W., and Wolever, T.M.S., 2008. Glycemic index of common Malaysian fruits. Asia Pacific
Journal of Clinical Nutrition 17:35-39.
Rocculi, P., Cocci, E., Romani, S., Sacchetti, G., and Rosa, M.d., 2009. Effect of 1-MCP treatment and N2O MAP on physiological
and quality changes of fresh-cut pineapple. Postharvest Biology and Technology 51:371-377.
Romero-González, R., Garrido Frenich, A., and MartÃnez Vidal, J.L., 2008. Multiresidue method for fast determination of
pesticides in fruit juices by ultra performance liquid chromatography coupled to tandem mass spectrometry. Talanta
76:211-225.
Ruma, G., Chakraborty, J., and Ghosh, D., 2008. Peroxidase activity of two cultivars (Kew and Queen) of ripe pineapple (Ananas
comosus) of Tripura. Journal of Applied Bioscience 34:106-109.
Sabbag, O.J., 2008. Environmental impact assessment post Euregap certification for pineapple production in Guaracai, Sao Paulo
State, Brazil. Pesquisa Agropecuaria Tropical 38:284-289.
Salomaoa, B.C.M., Slongo, A.P., and Aragao, G.M.F., 2007. Heat resistance of Neosartorya fischeri in various juices. LWT - Food
Science and Technology 40:676-680.
Sangsuwan, J., Rattanapanone, N., and Rachtanapun, P., 2008. Effect of chitosan/methyl cellulose films on microbial and quality
characteristics of fresh-cut cantaloupe and pineapple. Postharvest Biology and Technology 49:403-410.
Santos, M.d.D.M., Ribeiro, D.G., and Torres, A.C., 2008. Adventitious shoot of ornamental pineapple under benzylaminopurine,
naphthalene acetic acid and subculture period effect. / Brotações adventÃ-cias de abacaxizeiro ornamental sob o efeito de
benzilaminopurina, ácido naftalenoacético e perÃ-odos de subcultivo. Pesquisa Agropecuária Brasileira 43:1115-1120.
Santos, M.D.M., Buso, G.C.S., and Torres, A.C., 2008. Evaluation of genetic variability in micropropagated propagules of
ornamental pineapple [Ananas comosus var. bracteatus (Lindley) Coppens and Leal] using RAPD markers. Genetics and
Molecular Research 7:1097-1105.
Sarang, S., Sastry, S.K., and Knipe, L., 2008. Electrical conductivity of fruits and meats during ohmic heating. Journal of Food
Engineering 87:351-356.
Scordino, M., Sabatino, L., Gargano, M., Traulo, P., Pantó, V., and Gagliano, G., 2008. Assessment of fruit juice authenticity with
allergenic ingredients occurrence. / Valutazione dell'autenticità di succhi di frutta: con ingredienti allergenici. Ingredienti
Alimentari 7:14-18.
Sengupta, J., Ghosh, R., and Ghosh, D., 2008. Biochemical constituents of two cultivars of ripe pineapple (Ananas comosus) of
Tripura. Journal of Applied Bioscience 34:91-93.
Shamsudin, R., Daud, W.R.W., Takrif, M.S., Hassan, O., and Ilicali, C., 2009. Rheological properties of Josapine pineapple juice
at different stages of maturity. International Journal of Food Science & Technology 44:757-762.
Shen, B.N., Zheng, Y.X., Chen, W.H., Chang, T.Y., Ku, H.M., and Jan, F.J., 2009. Occurrence and molecular characterization of
three pineapple mealybug wilt-associated viruses in pineapple in Taiwan. Plant Disease 93:196-197.
Shi, X., Song, H., Huang, Z., Gao, T., and Xie, J., 2008. Research on the potential of fermentative hydrogen production from
pineapple peels at room temperature. Kezaisheng Nengyuan / Renewable Energy Resources 26:41-42, 45.
Siebeneichler, S.C., Monnerat, P.H., Carvalho, A.J.C.d., and Silva, J.A.d., 2008. Boro em abacaxizeiro 'Perola' no norte
fluminense-teores, distribuicao e caracteristicas do fruto (Boron in pinapple plants 'Perola' in the north fluminense-contents,
distribution and characteristics of the fruit). Revista Brasileira de Fruticultura 30:787-793.
Siebeneichler, S.C., Monnerat, P.H., and da Silva, J.A., 2008. Boron deficiency in pineapple 'Perola'. Acta Amazonica 38:651-656.
Silva, A.B.d., Pasqual, M., Castro, E.M.d., Miyata, L.Y., Melo, L.A.d., and Braga, F.T., 2008. Natural light in pineapple (Ananas
comosus L) micropropagation. / Luz natural na micropropagação do abacaxizeiro (Ananas comosus L. Merr). Interciencia
33:839-843.
Silva, A.E.d., Kadir, M.A., Aziz, M.A., and Kadzimin, S., 2008. Callus induction in pineapple (Ananas comosus L.) cv. Moris and
Josapine. International Journal of Agricultural Research 3:261-267.
Silva, J.M.d., Silva, J.P., and Spoto, M.H.F., 2008. Physico-chemical characteristics of pineapple submitted to ionizing radiation
technology as a method of post-harvest conservation. Ciência e Tecnologia de Alimentos 28:139-145.
Silva, J.S., Andreo, M.A., Tubaldini, F.R., Varanda, E.A., Rocha, L.R.M., Brito, A.R.M.S., Vilegas, W., and Hiruma-Lima, C.A.,
2008. Differences in gastroprotective and mutagenic actions between polar and apolar extracts of Ananas ananassoides.
Journal of Medicinal Food 11:160-168.
Silva, W.A., Carvalho, D.F., Varella, C.A.A., and Ceddia, M.B., 2008. Algorithm to mapping net income applied in irrigated
agriculture planning. / Sistema de informação geográfica para mapeamento da renda lÃ-quida aplicado no planejamento
da agricultura irrigada. Engenharia AgrÃ-cola 28:76-85.
Singh, C., Sharma, H.K., and Sarkar, B.C., 2008. Optimization of process conditions during osmotic dehydration of fresh
pineapple. Journal of Food Science and Technology 45:312-316.
Singh, S.K., Sharma, H.P., Jaivir, S., Vivak, K., and Special, 2007. Evaluation of physico-chemical and sensory qualities of mixed
fruit squash. Environment and Ecology 25S:1040-1045.
Singh, S.P., Tarsikka, P.S., and Harminder, S., 2008. Study on viscosity and electrical conductivity of fruit juices. Journal of Food
Science and Technology (Mysore) 45:371-372.
Slongo, A.P. and Aragão, G.M.F.d., 2007. Thermal resistance evaluation of Byssochlamys nivea and Beosartorya fischeri in
pineapple juice. Boletim do Centro de Pesquisa e Processamento de Alimentos 25:217-224.
Sopade, P.A., Halley, P.J., Cichero, J.A.Y., Ward, L.C., Liu, J., and Varliveli, S., 2008. Rheological characterization of food
thickeners marketed in Australia in various media for the management of dysphagia. III. Fruit juice as a dispersing medium.
Journal of Food Engineering 86:604-615.
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Newsletter of the Pineapple Working Group, International Society for Horticultural Science
Souza, C.B.d., Silva, B.B.d., Azevedo, P.V.d., and Silva, V.d.P.R.d., 2008. Energy fluxes and development of pineapple crop. /
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Stirling, G.R. and Pattison, A.B., 2008. Beyond chemical dependency for managing plant-parasitic nematodes: examples from the
banana, pineapple and vegetable industries of tropical and subtropical Australia. Australasian Plant Pathology 37:254-267.
Syue, P., Tang, C., and Lee, T., 2008. Studies on fruits internal browning of pineapple (Ananas comosus (L.) Merr.) by harvest
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Tamaki, V., Mercier, H., and Nievola, C.C., 2007. In vitro cultivation of clones of Ananas comosus (L.) Merril cultivar 'Smooth
Cayenne' using different macronutrients concentrations. Hoehnea 34:67-73.
Tancharoensukjit, S. and Chantanawarangoon, S. 2008. Effect of controlled atmosphere on quality of fresh-cut pineapple cv.
Phuket, Bangkok; Thailand.
Telteboim, M.C., Miranda, S.H.G.d., Oliveira, L., and Ozaki, V.A., 2007. Maximum residue limits and its implications on
international trade in fruits. Revista de Politica Agricola 16:102-112.
Thangaselvabai, T., Joshua, J.P., Justin, C.G.L., and Jayasekhar, M., 2007. Productivity and profitability of spice based cropping
system under forest ecosystem of Kanyakumari district. Journal of Plantation Crops 35:178-180.
Theerakulkait, C. and Sukhonthara, S. 2008. Effect of pineapple juice, pineapple shell extract and rice bran extract on browning in
banana [Musa (AAA Group) 'Gros Michel'] slices and puree, Bangkok; Thailand.
Theng, V., Agarie, S., and Nose, A., 2007. Regulatory properties of phosphoenolpyruvate carboxylase in crassulacean acid
metabolism plants: Diurnal changes in phosphorylation state and regulation of gene expression. Plant Production Science
10:171-181.
Theng, V., Agarie, S., and Nose, A., 2008. Regulatory phosphorylation of phosphoenolpyruvate carboxylase in the leaves of
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Tofanelli, M.B.D., Fernandes, M.d.S., Carrijo, N.S., and Martins Filho, O.B., 2008. Fresh fruits and vegetables market in Mineiros,
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Fruits:23-24.
Zhang, Y.-Y., Hsu, H.-J., and Huang, W.-L., 2008. Establishment of callus induction and shoot regeneration system from axillary
bud on Taiwanese edible pineapples. Taiwanese Journal of Agricultural Chemistry and Food Science 46:49-56.
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Chinese Journal of Animal Nutrition 20:645-650.—
Instructions to Contributors to Pineapple News
All contributions should be written in English. Editing assistance will be provided on request.
Preferred contributions include:
•
Tim ely news about research on issues related to culture, processing, storage, and m arketing of pineapple.
•
New, interesting, or unique problem s encountered by growers.
•
Country or status reports on the local pineapple industry.
•
If uncertain about the suitability of m aterial for the newsletter, contact the editor.
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If possible, please send contributions by E-m ail as attached files in MS W ord or rich text form at or on floppy disks.
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Pineapple News is available on the W eb at: http://tpss.hawaii.edu/pineapple/pineappl.htm .—
Pineapple News is published by the University of Hawaii, College of Tropical Agriculture and Human Resources, Dept. of Tropical
Plant and Soil Science. The contributions of Valent Biosciences and by various individuals to the University of Hawaii Foundation
help support the costs associated with the publication of the newsletter. Reference to commercial products and services is made
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The University of Hawaii, College of Tropical Agriculture and Human Resources is an Equal Opportunity and Affirmative Action
Employer.
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