AN ABSTRACT OF THE THESIS OF
for the
Herman Vaartnou
(Name)
Farm Crops
in
presented
Doctor of Philosophy
(Degree)
on December 16, 1966
(Major)
Title:
(Date)
RESPONSES OF FIVE GENOTYPES OF AGROSTIS L.
TO VARIATIONS IN ENVIRONMENT
Abstract approved:
Redacted for Privacy
(Signature)
Morphological changes in five clonally propagated genotypes
of
Agrostis L. resulting from variations in environment were
studied in six natural environments extending from southern Oregon
to northern British Columbia.
These same clones were also sub-
jected to four different cutting heights in turf plots. In addition to
the field experiments, they were also studied in growth chambers
using four different combinations of day and night temperatures.
Tiller number, plant diameter, leaf width, leaf length, aerial
branching, rhizome number, rhizome length, nodes per rhizome,
stolon number, stolon length and the number of stolon nodes were
the morphological characteristics studied.
Consistent morphological differences indicated that the five
clones do not belong to the same species. Detailed taxonomic
study of the population from which these clones were selected is
warranted.
The five clones varied in their capacity to
of
tiller. The density
tillers depended both upon environment during the time
of
tiller
development and clonal genetic differences. Experimental genotypes
differed greatly in their susceptibility to Fusarium sp.
,
therefore
causing some of the differences in tiller density during the spring
observation time.
The genotypes reacted differently to cutting
height defoliation during different seasons.
They also reacted
differently to temperature, day length and light intensity.
The five genotypes each had different leaf widths and leaf
lengths when grown in similar environments.
Increased light in-
tensity and day length increased the leaf width at locations where
the temperature had not affected the development of leaves and
initiation of leaf primordia. Within one temperature combination
one genotype produced the
narrowest leaves and the widest leaves
were produced by the other four. The length of leaves depended
mainly upon the genotype.
No
correlation between leaf width and
length was observed.
Aerial branching occurred in times when the dominance of
the apical bud was removed and the lateral buds developed.
Cor-
relation between tillering and aerial branching within clones was
observed.
Clonal differences accounted for most of the variation in
rhizome number. Rhizome length was influenced by clones,
environment and the clone x environment interaction.
No
correla-
tion existed between the clones ability to form tillers and rhizomes.
The clones formed most stolons in the fall after the
sterile
shoots had been formed. Short days and lower light intensity in
the fall combined with low night temperatures were the environ-
mental factors which promoted the formation of stolons.
The
ability of clones to form stolons was not related to their ability to
form rhizomes or tillers.
Responses of Five Genotypes of Agrostis L.
to Variations in Environment
by
Herman Vaartnou
A THESIS
submitted to
Oregon State University
in partial fulfillment of
the requirements for the
degree of
Doctor of Philosophy
June 1967
APPROVED:
Redacted for Privacy
Associate Professor ?of Farm Crops
in charge of major
Redacted for Privacy
Head a Department of Farm Crops
Redacted for Privacy
Dean of Graduate School
Date thesis is presented
Typed by Kay Smith for
December 16, 1966
Herman Vaartnnu
ACKNOWLEDGMENT
Sincere thanks are extended to Dr. N. R. Goetze and Dr,
V. C.
Brink for their assistance and encouragement throughout the study
and for their helpful
criticism
of the manuscript.
Most sincere thanks is extended to Mr. Leon J. Koerner for
his encouragement in times when it was most needed.
Appreciation is expressed to Dr.
D. O.
Chilcote and Dr. K. L.
Chambers for assistance in reviewing the manuscript and
Guitard, Dr. J. Miltimore, Mr.
W.
to Dr. A. A.
Burnes, Mr. J. Yungen and
Mr. A. E. Gross for providing the land for experiments and main-
taining the plants during the study period.
Thanks are also due to Mr. C. D. Davenport and Mr. M,
Vaartnou for assistance in the preparation of the manuscript,
Last but not least,
I
am grateful to The Leon and Thee Koerner
Foundation for financial help.
To my wife Hella and sons
Manivalde, Peter and Erik
TABLE OF CONTENTS
Page
I.
II.
Introduction
1
Literature Review
3
Genotypes
3
Tillering
5
9
Root Growth
Growth of Rhizomes
Growth of Leaves
III.
Materials and Methods
Source and Description of Clones
Clone A
Clone B
Clone C
Clone D
Clone E
Field Experiment with Single Spaced Plants
Environments
Clonal Increase
Planting and Maintenance
Observation Times
Types of Observation
Turf Experiment
Growth Chamber Experiment
IV.
Results
Tillering
Field Experiments with Single Spaced Plants
Growth Chamber Experiment
Turf Experiment
Diameter of Plant
Field Experiment with Single Spaced Plants
Growth Chamber Experiment
Growth of Leaves
Field Experiment with Single Spaced Plants
Growth Chamber Experiment
11
12
15
15
15
20
26
31
35
39
39
44
48
49
49
51
54
56
56
56
68
73
75
75
78
79
79
89
Page
Aerial Branching
Field Experiment with Single Spaced Plants
Turf Experiment
Growth of Rhizomes
Field Experiment with Single Spaced Plants
Growth Chamber Experiment
Turf Experiment
Growth of Stolons
Field Experiment with Single Spaced Plants
Turf Experiment
V.
Discussion
Tillering
Growth of Leaves
Aerial Branching
Growth of Rhizomes
Growth of Stolons
VI.
91
91
91
95
95
103
103
108
108
112
119
119
122
124
125
127
Summary
128
Bibliography
131
Appendix
138
LIST OF TABLES
Page
Table
1
2
3
4
5
6
7
8
9
10
The monthly temperature and rainfall data at
Medford Experimental Station in the years
1964 and 1965.
41
temperature and rainfall data at
Klamath Falls Experimental Station in the
years 1964 and 1965.
42
The monthly temperature and rainfall data at
Vancouver, B. C. in the years 1964 and 1965.
43
The monthly
temperature and rainfall data at
Summerland Experimental Station in the years
The monthly
1964 and 1965.
45
The monthly temperature and rainfall data at
Smithers Experimental Station in the years
1964 and 1965.
46
temperature and rainfall data at
Beaverlodge Experimental Station in the years
The monthly
1964 and 1965.
47
Average number of basal tillers per 25 square
centimeters at three dates for five clones of
Agrostis L. grown at six locations.
57
Average number of basal tillers per plant and
average diameter of plant for five clones of
Agrostis L. grown at four different day and
night temperatures.
69
Average number of tillers per cm. 2 for five
clones of Agrostis L. grown under four
different cutting heights.
74
Average diameter of plant at three dates for
five clones of Agrostis L. grown at six locations. 76
Page
Table
11
12
13
14
15
16
17
18
19
20
21
Average leaf width at three dates for five clones
of Agrostis L. grown at six locations.
80
Average leaf length at three dates for five clones
of Agrostis L. grown at six locations.
86
Average leaf width and leaf length for five clones
of Agrostis L. grown at four different day and
night temperatures in combinations.
90
Average number of aerial branches at three dates
for five clones of Agrostis L. grown at six locations.
92
Average number of aerial branches per one
hundred cm.2 area of five clones of Agrostis L.
grown under four different cutting heights and
observed at three times.
94
Average number of rhizomes per plant at three
dates for five clones of Agrostis L. grown at
six locations.
96
Average length of rhizomes at three dates for
five clones of Agrostis L. grown at six locations.
99
Average number of rhizome nodes at three dates
for five clones of Agrostis L. grown at six
locations.
101
Average number of rhizomes per plant, average
length of rhizomes and average number of
rhizome nodes per rhizome for five clones of
Agrostis L, grown at four different day and
night temperatures.
104
Average weight of rhizomes per 100 cm.2 at
three dates for five clones of Agrostis L.
grown under four different cutting heights.
105
Average length of rhizomes at three dates for
five clones of Agrostis L. grown under four
different cutting heights.
106
Page
Table
21
22
23
24
25
26
27
28
Average length of rhizome at three dates for
five clones of Agrostis L. grown under four
different cutting heights.
106
Average number of rhizome nodes per rhizome
at three dates for five clones of Agrostis L.
grown under four different cutting heights.
107
Average number of stolons at three dates for
five clones of Agrostis L. grown at six locations.
109
Average length of stolons at three dates for five
clones of Agrostis L. grown at six locations.
110
Average number of stolon nodes at three dates
for five clones of Agrostis L. grown at six
locations
113
Average weight of stolons per 100 cm. 2 area
at three dates for five clones of Agrostis L.
grown under four different cutting heights.
115
Average length of stolons at three dates for
five clones of Agrostis L. grown under four
different cutting heights.
117
Average number of stolon nodes per stolon at
three dates for five clones of Agrostis L. grown
under four different cutting heights.
118
Mean squares and levels of significance for
number of tillers, leaf width, leaf length and
diameter of plant. Observation time as a
main plot.
138
Mean squares and levels of significance for
number of rhizomes, length of rhizomes and
number of rhizome nodes. Observation time
as a main plot.
140
Appendix
Table
1
2
Appendix
Table
3
4
5
6
7
8
9
10
Page
Mean squares and levels of significance for
number of stolons, length of stolons, number
of stolon nodes and number of aerial branches.
Observation time as a main plot.
Mean squares and levels of significance for
number of tillers, leaf width, leaf length and
diameter of plant. Location as a main plot.
Mean squares and levels of significance for
number of rhizomes, length of rhizomes and
number of rhizome nodes. Location as a
main plot.
Mean squares and levels of significance for
number of stolons, length of stolon, number
of stolon nodes and number of aerial branches.
Location as a main plot.
142
144
145
146
Mean squares and levels of significance for
number of tillers, amount of rhizomes, length
of rhizomes and number of rhizome nodes. Turf
experiment.
147
Mean squares and levels of significance for
number of aerial branches, amount of stolons,
length of stolons, and number of stolon nodes.
Turf experiment.
148
Mean squares and levels of significance for number of tillers, leaf width, leaf length and diameter
149
of plant. Growth chamber experiment.
Mean squares and levels of significance for
number of rhizomes, length of rhizomes, and
number of rhizome nodes. Growth chamber
experiment.
149
LIST OF FIGURES
Page
Figure
1
Clone A growing at Vancouver, B. C.
2
Pattern of rhizome and tiller growth of clone
17
A
grown at Vancouver, B. C.
17
3
Clone A grown at Summerland, B. C.
18
4
Panicles from clones grown at Vancouver,
B. C.
18
5
Clone A growing at Vancouver, B. C.
19
6
Ligule of clone A.
19
7
First glume
21
8
Lemma, palea, and grain of clone A.
21
9
Clone
B
growing at Vancouver, B. C.
23
10
Clone
B
growing at Smithers, B. C.
23
11
Clone
B
grown at Summerland, B. C.
24
12
Clone
B
growing at Vancouver, B. C.
24
13
Ligule of clone B.
25
14
First glume
25
15
Lemma, palea, and grain of clone B.
27
16
Clone
growing at Vancouver, B. C.
27
17
Pattern of rhizome and tiller growth
clone
18
C
C
of clone A.
of clone B.
grown at Vancouver, B. C.
Ligule of clone C.
of
29
29
Page
Figure
19
First glume
20
Lemma, palea, and grain of clone C.
21
Clone
22
Ligule of clone D.
33
23
First glume
34
24
Lemma, palea, and grain of clone
D.
34
25
Clone E growing at Vancouver, B. C.
36
26
First glume
of clone E.
36
27
Lemma, palea, and grain of clone E.
38
28
Clone A growing at Beaverlodge, Alberta.
59
29
Clone
growing at Beaverlodge, Alberta.
59
30
Clone A growing at Smithers, B. C.
60
31
Plants grown at Summerland,
B. C.
60
32
Plants grown at Summerland,
B. C.
62
33
Average number of tillers of five clones of
Agrostis L. grown at Beaverlodge and Smithers
and measured at three different observation
times.
65
Average number of tillers of five clones of
Agrostis L. grown at Summerland and
Vancouver and measured at three different
observation times.
66
Average number of tillers of five clones of
Agrostis L. grown at Klamath Falls and
Medford and measured at three different
observation times.
67
34
35
B
C
of clone C.
growing at Vancouver,
30
B. C.
of clone D.
30
33
Figure
Page
Average number of basal tillers per plant of
five clones of Agrostis L. grown 60 days at
four different day and night temperatures
in the growth chamber.
70
Clone D grown 60 days in the growth chamber
at 60 °F. day and 45 °F. night temperatures.
71
Clone A grown 60 days in the growth chamber
at 60 °F. day and 45 °F. night temperatures.
71
39
Plants grown
60
days in the growth chamber.
72
40
Plants grown
60
days in the growth chamber.
72
41
Average leaf width of five clones of Agrostis
L. grown at Beaverlodge and Smithers and
measured at three different observation times.
81
Average leaf width of five clones of Agrostis
L. grown at Summerland and Vancouver and
measured at three different observation times.
83
Average leaf width of five clones of Agrostis
L. grown at Klamath Falls and Medford and
measured at three different observation times,
84
36
37
38
42
43
RESPONSES OF FIVE GENOTYPES OF AGROSTIS L.
TO VARIATIONS IN ENVIRONMENT
INTRODUCTION
Bentgrasses, Agrostis
L.
,
are among the most important
turfgrass species in the Pacific Northwest. Besides being
of un-
matched value for turf purposes, they have gained importance as
an income source for seed growers.
Although this grass can be
found in every garden and park in the Pacific Northwest, very little
research work
on its management has been
onomic study exists as yet.
carried out.
No tax-
Only limited breeding studies on the
species Agrostis tenuis Sibth. and Agrostis gigantea Roth have been
made.
Although these two species provide most of the material for
permanent turf areas, few articles have been published on the ecology of the genus within the
last fifty years. The latitudinal adapta-
tions of bentgrasses have not been determined. Also unknown at
present is whether day temperature, night temperature, or disease
limit the area suitable for bentgrasses.
To
clarify some of the points mentioned previously, this study
was initiated and carried out in six widely separated natural environ-
ments ranging from southern Oregon to northern British Columbia,
using five vastly different clonally propagated genotypes of bent -
grasses.
2
The same clones were tested as single plant cultures under
controlled light and different temperature conditions in growth
chambers. Their reaction to different cutting heights in turf plots
under field conditions were also studied.
3
LITERATURE REVIEW
Genotypes
Genetic and physiological factors determine the morphological
features of
a plant growing under
natural environment. The ultimate
limit would be set by genetic factors and within those limits the
physiological factors would determine the morphological features (65)
under the environment of the experiment.
Local areas, charac-
terized by differences in substrate, topography or microclimate,
can produce clearly recognizable races within a species (27). Alti-
tudinal and latitudinal climatic gradients are well known for their
ability to produce ecological races (27, 48). Often we can find, very
shortly after introduction of a species into a new area, two genetically distinct ecotypes evolving (11).
We cannot be
sure if these
ecotypes evolved before their introduction or whether they were
selected after their introduction by the new environment. Working
with Agrostis tenuis, Jowett (28) suggested that rapid evolution can
occur with little spatial isolation, and that adaptive significance is
indicated by changes in morphological characters.
Philipson (49) describes Agrostis tenuis as a perennial with
short rhizomes and occasional stolons.
above and closed near the base.
blunt.
The leaf sheaths are open
The ligule is short, broad and
Panicle branches and branchlets remain open in fruiting.
4
It has a radical leaf with a linear, pale to dark green blade.
describes Agrostis gigantea as
a
perennial with spreading rhizomes
Leaf sheaths split to near the
and stolons, forming an open tuft.
The ligule is longer than broad and is rounded.
base.
He
In
fruiting,
the main branches remain spreading and the branchlets close against
them. It has a radical leaf with the blades, usually gray -green or
green, broadest above the base.
Bjorkman (8), working with plants of Agrostis tenuis from
natural habitats, found the chromosome number to
with a few trisomics and some 2n
=
28
+
be
largely 2n
=
28
ss. Stuckey and Banfield
(62), however, reported finding nearly all numbers between 28 and
42 in
bentgrass plants raised from seed collected in nature. They
found the progeny of such plants to exhibit great variations, both
morphologically and in the chromosome number. There was
no
evidence of any correlation between chromosome number and plant
morphology.
Carrol (10), working with'Highland' and Agrostis tenuis plants,
found that 'Highland' was injured more by low soil temperature than
Agrostis tenuis.
He found
than air temperature.
that soil temperature was more injurious
The lethal minimum soil
temperature was
between -10 degrees to -15 degrees Centigrade and the lethal maximum soil temperature was between
grade.
50
degrees to
60
degrees Centi-
Plants from high nitrogen plots suffered more than plants
5
from low nitrogen plots. Hiesey (26) found that Poa L. species
responded best to different day and night temperatures similar to
the seasonal temperatures at their natural habitat.
Beard (5) found that the micro - environment in shade promotes
disease activity, which is more important in affecting the adaptation
of
turf species to shade than light, moisture or nutrient deficiencies.
Juska (29), using eleven bentgrass varieties, concluded that turf
grown under shade conditions would be lighter green, less dense and
more succulent.
He
hypothesized that the relation between varieties
would be the same in shade as in sunlight whenever disease preven-
tion was practiced.
Tillering
Evans and Ely (17), working with Phalaris arundinacea L.
,
pointed out that aboveground shoots develop mainly in fall and early
spring.
The shoots developing in spring normally completed their
life cycle in the same year.
The shoots developing in fall completed
their life cycle the following year.
Environmental factors affect both the rate of leaf production
and tillering since the new
leaves (69).
tillers arise from buds
in the axils of the
Langer (35) showed that the number of tillers per unit
area declined from early spring to the middle
of
summer, and
suggested that this phenomenon is quite common in grasses.
6
Silsbury (55) studied Lolium perenne and indicated that not more
than
10
percent of the vegetative tillers present in the early summer
are able to regrow in the following autumn. Auxiliary buds provide
the new tillers and regrowth in the fall.
Bromus inermis L.
,
Lamp (34), working with
found that most of the
tillers emerging
in spring
failed to survive and that they do not contribute greatly to the total
development of the plant.
The number of bud
of a plant.
primordia determines the tillering capacity
It is an hereditary
the species or clone (14, 41).
characteristic and varies according to
Growth of the lateral buds can be
promoted by reducing the daylength or light intensity, high tempera-
ture or by partial defoliation.
The
importance of
single factor
a
depends on the level of other factors (41). Environmental effects on
tillering make it nearly impossible to compare tillering for one year
with that of another (59).
Alberda (1), working with Lolium perenne L.
,
found that
cutting provides an initial decline in tillering. When tillering re-
commences, it occurs more vigorously than previously.
Removing
the growing points through cutting promoted tiller production in
Italian ryegrass, Lolium multiflorum Lam. (39). However, suc-
cessive cropping reduced the number of tillers in barley, Hordeum
vulgare
L. (21).
Zavitz (73), working with oats, Avena sativa L.
,
found that
7
vigorously tillering plants are able to adjust better to different
cultural and environmental conditions than light tillering ones.
Increasing the soil temperature from
22
degrees Centigrade
to 42 degrees Centigrade reduced the number of
wheat, Triticum vulgare Host. (71).
tillers
in Marquis
Lowering the temperature or
raising the light intensity increased the tillering of ryegrass, Lolium
L. (40).
When the number of individual
a wide range of
tillers showed little change,
temperatures existed. Rapid reduction
numbers occurred above
85
in
degrees Fahrenheit and below
tiller
55
degrees
Fahrenheit (42). Lowering the night temperature increased the
tillering in several grasses (43). Mitchell and Lucanus (44), working
with browntop, Agrostis tenuis, and yorkshire fog, Holcus lanatus
stimulated tillering by low
Low light
Pers.
L.
40
,
degrees Fahrenheit night temperatures.
intensity delayed the tillering of Sorghum vulgare
(53) and also
L. (15).
35
decreased tillering number in Bromus mollis
Bean (4), working with 'S -37' cocksfoot, Dactylis glomerata
found that when light intensity was decreased to 60 percent and
percent of full daylight, the rate of tiller production decreased.
Nittler, Kenny and Osborne (46) found that varietal differences in
tiller production were small when light was available
in sufficient
quantities, but very large when light was a limiting factor.
Evans (16), studying the life history of timothy, Phleum
pratense
L.,
L.
,
found that the age of the plant was important in
8
tillering. Young seedlings tillered best in fall with short photo periods. Plants in an established meadow can tiller any time, but
do so in
greatest numbers about midsummer. Other environmental
factors, such as temperature, light intensity, moisture, nutrition
and cutting, modified the effect of daylength.
The length of day
factor can profoundly alter the vegetative growth in herbaceous
grasses (2).
It also leads to
suppression of the reproductive develop-
ment and the dominance of vegetative growth resulting in vigorous
sod formation by increased tillering.
Using southern grasses as
experimental material, Knight and Benedict (33) found that tillering
increased when daylength decreased below
14
hours, and especially
when night temperature fell below 55 degrees Fahrenheit.
Plants of
Bromus inermis produced rosettes when grown during short days of
8.
5
hours. Under these conditions the number of tillers was great-
est (66).
For tillering there are different optimum soil moisture contents
for different species and varieties of grasses (56, 47).
to Leopold (36),
According
tillering in barley can be controlled by auxin, and
in alfalfa, Medicago sativa
L., by using antiauxin (13). Apparently
nitrogen is the limiting factor in tillering (12, 21, 31) and together
with different temperature combinations (52), can control the number
of
tillers formed
Poa pratensis L.
on
,
perennial grasses.
Darrow (14), working with
reports that while using ammonium nitrogen,
9
the best growth was obtained at pH 6. 5, but if nitrate were used as
source of nitrogen, any pH between 4.
a
the same result.
5
and 6.
5
produces about
Normally, mineral nutrition deficiency reduced
the number of tillers (21, 53, 67).
.
However, it may be that the
balance between the supply and consumption of different mineral
elements plays a role in controlling the number of tillers per plant
(35).
Brenchley (9), working with barley, and Grantham (22), work-
ing with wheat, reported on the importance of phosphorus at certain
developmental phases of the plants.
decreased the number of tillers.
on
A
deficiency of phosphorus
Potassium had very little effect
tillering under the experimental conditions (22). Boron deficiency
increased the number
of
tillers
in wheat (45) and in
barley (64).
Knight (32) did not associate the decline in tillering in Dactylis
glomerata with the onset of the reproductive phase.
Root Growth
Hanson and Juska (24) found that the roots of Merion Kentucky
bluegrass, Poa pratensis
L.
September and April.
least one -half
At
,
make most of their growth between
bluegrass (58) developed in springtime.
of the
roots of Kentucky
This is in agreement with
Stuckey's (60) findings on bentgrass and fescues, Festuca L., where
most of the old roots disintegrated after the new ones developed.
However, the report disagrees with conditions found by Stuckey (60)
10
existing in Kentucky bluegrass where most of the roots were of
perennial nature and only
a few new ones developed each
spring.
Frequent cutting normally reduces the root growth much more
than it reduces the top growth (37, 38).
It is possible (60) that the
high soil temperature in summer inhibits root growth. According
to Stuckey (60), roots are able to grow at
degrees Fahrenheit. Beard and Daniel
temperatures close to
(6)
32
blame poor aeration, high
moisture, and high soil temperature for the midsummer dieback of
roots of bentgrass turf.
from
60
They found that increasing the temperature
degrees Fahrenheit to
90
degrees Fahrenheit decreased the
formation of new roots.
Soper (57) emphasized that high temperature combined with
low light intensity reduces the root growth of perennial
He pointed out
ryegrass.
that root growth is reduced more than shoot growth.
Working with three different temperatures, Stuckey (61) found that
bentgrass roots remained relatively unbranched and immature when
the experimental temperature was 50 degrees Fahrenheit.
soil temperatures of
60 and 80
Higher
degrees Fahrenheit resulted in early
root maturation and ultimate disintegration.
Youngner (72), working with Zoysia Willd.
,
found that maximum
root growth occurred at high temperatures (27 degrees Centigrade)
if these were accompanied by 14 to 16 hour photoperiods.
(14)
Darrow's
results show that Kentucky bluegrass roots, when grown under
11
low (15 degrees Centigrade) temperature, were succulent, white
At the same time they branched heavily.
and large.
These can be
compared with light brown, densely tufted, and small diameter roots
grown under high (35 degrees Centigrade) temperatures.
Excessive respiration, due to high temperatures, can lead to
the exhaustion of carbohydrate reserves in roots, and combined with
frequent cutting, to the death of the plants (63). Stimulating metabolic activity through nitrogen fertilization, defoliation or high tem-
peratures would have the same effect
on promoting or killing the
roots. Environmental factors, such as nutrients, moisture, soil
texture, temperature, and light, contribute to the balance of storage
reserves
in underground organs and general health of the plants (68).
Growth of Rhizomes
Kershaw (30) separated the rhizomes of Agrostis tenuis into
two categories: (a) pioneer rhizomes
-
normally situated at a deep
level in the ground and not often found tillering, and (b) colonizing
rhizomes
a
-
these developing much closer to the surface and forming
tiller or an aggregate
of
tillers.
The aggregation of
separate rhizome systems is proposed as
a
tillers from
basic feature for the
cyclic phases in vegetation and is suggested as a widespread phenomenon in rhizomatous species.
Hansen and Juska (24) found that there is very little increase
12
in rhizome growth in Kentucky bluegrass from September to April.
In Kentucky bluegrass, rhizomes developed mainly during late spring
and the number per plant was much greater under the long daylength
season than under the short daylength season.
Contrary to the
response in Kentucky bluegrass, Canada bluegrass, Poa compressa
L.
,
developed the largest number of rhizomes in late fall or early
spring when the daylength was short (18).
Watkins (66), working with Bromus inermis, found that long
daylengths (15 hours) produced the maximum number of rhizomes,
but maximum size and length of rhizomes were produced in 18 hour
daylengths.
Ammonium sulphate was inferior to calcium nitrate in promoting the growth of rhizomes (25).
of nitrogen, a pH of between 4.
growth but a pH of 6.
5
5
Using calcium nitrate as the source
to 6.
5
did not affect the rhizome
was best for rhizome growth if ammonium
sulphate was used as the nitrogen source (14).
Wood and Burke (70) did not find any close association between
the ability to form a dense turf and rhizomes when they were working
with 'Merion', 'Park', 'Delta', and 'Newport' varieties of Kentucky
bluegrass.
Growth of Leaves
Ryder (50) suggests that leaf shape depends on heredity and it
13
may be changed by environment.
Ashby (3) suggests that leaf shape can be influenced by light
intensity, operating through carbohydrate metabolism, and by
mineral nutrition. Beinhart
(7) found
that light intensity affected the
activation of new meristems but not the rate at which leaves were
produced by already active meristems.
to produce large leaf
In
Low light
intensities tend
areas and to decrease the leaf thickness (54).
'Marquis' wheat (19), each increase in light intensity over the
range 200 to 2500 ft. c. resulted in an increase in breadth and thickness but
8
a
decrease in length
of leaf.
An
increase in daylength from
to 24 hours increased leaf length, breadth, and area.
It is sug-
gested that leaf growth is controlled by hormonal mechanisms which
are sensitive to photoperiod.
Competition for assimilates among
developing leaves does not seem to be the deciding factor.
Mitchell (42) found that the optimum temperature for the length
of the leaf blade would be 75
ryegrass, and
85
degrees Fahrenheit for browntop and
degrees Fahrenheit for Paspalum
L.
Ryle (52)
found that the optimum temperature for leaf width may be somewhat
lower than that for maximum leaf length in cocksfoot, ryegrass, and
fescue. High temperature increased the rate of leaf appearance,
the number of actively elongating leaves, and leaf length.
However,
the leaf width was reduced by high temperature.
If
environmental conditions delayed the appearance of the leaf,
14
the length of the initial leaves was reduced greatly (40).
working with winter rye, Secale cereale L.
the lamina length of the
,
Hansel (23),
and barley, found that
first and second leaf was decreased
by
ing the vernalizing temperature from -3 degrees Centigrade to
degrees Centigrade.
The extended period of vernalizing
rais5
further
shortened the lamina in winter grain but not in spring barley. Using
calcium nitrate on Kentucky bluegrass, Harrison (25) was able to
demonstrate a shortening of the leaf blade.
Nitrate nitrogen had a greater effect than ammonium nitrogen
on Kentucky bluegrass leaf development when the
was
15 to 35
degrees Centigrade and the pH was
temperature range
4. 5 to 6. 5 (14).
Some genotypes of Potentilla glandulosa L. can be recognized
only if grown under certain environmental conditions (11).
This
means that some of them can alter the expression of their phenotypic
character with
a change in
climatic condition.
Working with timothy, Ryle and Langer (51) found that in
elongated vegetative shoots, stolons may be produced when high
temperature or photoperiods inhibit the spikelet initiation.
Garner and Allard (20) demonstrated that changing the day length from optimum to sub - optimum increased the branching through
a
decrease in apical bud dominance which in turn led to activation of
the lateral buds.
15
MATERIALS AND METHODS
III.
Source and Description of Clones
The plant
material for this study was selected from various
old sod fields or turf installations in the Northern Pacific Coast
Region of North America between north latitudes of 45o and 55o.
For all of the five genotypes, single plant selections were made in
the Fall of 1963.
University of
B. C.
They were clonally propagated and grown in the
,
Division of Plant Science greenhouse during
the winter of 1963 -64.
Clone A
This was a clone from 'Highland' bentgrass which was collected
in 1963 by Dr. N. Goetze and Mr. H. Schoth, both of Oregon State
University, from an old 'Highland' bentgrass seed field belonging
to Mr. Don Savage of Silverton, Oregon.
to be 15 years old and is located in the
grass producing area
of Marion County.
This field was estimated
heart
of the 'Highland' bent -
Dr. Goetze and Mr. Schoth
spent considerable time selecting this clone as the most typical of
what is normally called 'Highland' bentgrass.
16
Morphological characteristics
Perennial; forming numerous, long branching rhizomes.
Plant.
Secondary rhizomes easily extend to surface and then
tiller slightly (Figure
40 - 80 cm. high,
Culm.
1,
2).
glabrous; sterile shoots forming
long trailing stolon in fall (Figure 3).
Fertile shoots
erect, sometimes slightly geniculate at base (Figure
Panicle.
10 - 18
1).
cm. long; pyramidal in form; reddish brown,
particularly the nodes
of the panicle.
Open in flower,
Branches open, branchlets
semiclosed at fruiting.
closed against them (Figure 4).
Main nearly smooth, branches scabrous and slightly
Rachis.
toothed.
Rachilla.
Leaf.
Not extending beyond the floret.
Sheath
split nearly to the base. Blade flat, usually
1. 0 - 2. 5
mm. wide, 50
- 70
mm. long, bluish green,
wider just above the base, tapering evenly towards the
tip (Figure 5).
Elongated, more length than breadth;
Ligule.
1
- 4
mm. long,
rounded and toothed, sometimes split (Figure 6).
Glume s
.
Outer
-
2.2
- 3. 2
mm. long,
0.6 -
0. 7
mm. wide.
Lanceolate, tapering evenly towards the tip.
Edges and
17
Figure
1.
Clone A growing at Vancouver, B. C.
Photographed August 20, 1965.
Figure
2.
Pattern of rhizome and tiller growth
of clone A grown at Vancouver, B. C.
Photographed September 1, 1965.
b.
Figure
3.
'
-,
18
`^3
Alp
Clone A grown at Summerland, B. C.
This clone produced only a few flowering
shoots because of winter damage. The
trailing stolons are characteristic of this
clone.
Figure 4.
Photographed September
5, 1965.
Panicles from clones grown at
Vancouver, B. C. From left to
right; clones B, A, C, E, D.
Photographed September 1, 1965.
19
a
-
-
;r
',,+-I
.......
w v_
....a
ti
(
""-,...
_.
r-'
_
Figure
eR_+
k
'
r
- F-_
;. 1 -.`
'
-
-.
..ü
F
"A.
5.
..
1
4
'
1
.
1}
1
'r
r. ld
.
o
,,
-,
w
`
Se,
-
aTEINnpm
-
nori,
,
.;
Clone A growing at Vancouver, B. C.
In the fall long, trailing sterile shoots
were formed. Photographed September
2, 1964.
;'
Figure
6.
Ligule of clone A.
20
center nerve finely toothed from middle to tip (Figure 7).
Inner
- 2.
1
- 2. 8
mm. long, 0.5
0.65 mm. wide, elliptic,
tapering evenly towards the ends. Edges and midnerve
toothed from center to tip.
Lemma.
Elliptic; silvery;
wide (Figure 8).
face.
1. 5 -
1.9 mm. long, 0.42 - 0.48 mm.
Slightly scabrous over the whole sur-
Covered with few unicellular pilose hairs.
Three
nerves reaching the apex of lemma, sometimes slightly
excurrent.
Callus. With two lateral fasicles of hairs. Hairs unicellular,
less than 1/4 of lemma.
Palea.
Two -nerved,
tapers evenly from the base toward the
two - lipped apex (Figure 8).
0. 35 mm. wide.
0.9
- 1. 2
mm. long, 0.25
-
Partially adhering to the grain; hyaline.
Grain. Free; enclosed in the floret; broadest at the middle;
depressed in front.
1. 2 - 1. 4
mm. long, 0. 4
-
0. 5 mm,
wide (Figure 8).
The morphological
characteristics
of this clone
are very
similar to the morphological characteristics of Agrostis gigantea
Roth. as described by Philipson (49).
Clone
B
This clone was collected in 1963 by Dr.
N.
Goetze, Oregon
21
,y.
--"'~°r
Figure
7.
First glume
of clone A.
1
.1
se-
Figure 8. Lemma, palea, and grain of clone
A.
22
State University, and Mr. J. Wood, Clatsop County Agent, from an
old abandoned, hand -dug dike on the Mr. Dave Hess farm near
Astoria, Oregon.
This area has not been disturbed since the late
The plant was most typical of what is called
19th century.
'Astoria'
bentgrass.
Morphological characteristics
Perennial; caespitose; sometimes forms
Plant.
spar-
a few,
ingly branching rhizomes (Figure 9).
Culms. 40
- 80
cm. high; glabrous.
Fertile shoots upright
or slightly geniculate at base (Figure 10).
In
favorable
environment the sterile shoots grow as stolons in the
fall (Figure 11).
Panicle.
12 - 20
cm. long; lanceolate; brown at maturity.
Open at fruiting (Figure 4).
Rachis. Main nearly smooth, branches scabrous and slightly
toothed.
Rachilla.
Not extending beyond the floret.
Sheath closed near the base, open above.
Leaf.
usually 2.
5 -
Blade
4. 0 mm. wide, 70 - 90 mm. long,
nearly
linear, light green, flat (Figure 12).
Ligule.
Less length than breadth;
1
- 2
mm. long; truncate.
Sometimes slightly elongated, split and toothed (Figure
13),
23
' V;;:;:ir
-"rTur,l':744,t
4!'
.r
N.
,
'
,
.
'
il "...4.4".''','-
;"1.>"'
'°(,:%-:?,:tz
4.
,..
Ar'' L '
l7'
I:-
,I4 +.;,,,../$7.4.
41, '-;
*:
14..1
..----
-
'
,
'.
-
.
V.'
'
,
N
.7.,....y
.
:
cr,..1"4
Figure
10.
NIT
.
9tiltik,\
,s1'
T: ntis
17T
Figure 9.
'W
.
.
'
'
:Le
'.3
4
growing at Vancouver,
Photographed August 20, 1965.
Clone
B
B. C.
Clone B growing at Smithers, B. C.
The clone is very vigorous. Photographed July 3, 1965.
24
,V
.q
f,,r
r. ~ áa
1
N.
s,
-.:4
:.
i
,
i
)
.,
1
F
\I
, /
'
`
I
1.
.
.
.
M: ,;,
V
1
_'
, -
=br
\
.
_
_`
f
IQ
.
Figure
11.
grown at Summerland, B. C.
This clone produced many fertile shoots
in spring and only a few stolons in the
Clone
B
fall.
Photographed September
*
P;,
5, 1965.
Pi..-,,
-
2
wr
.
:S
"
..
s-
a71--
f-'
i
7: v3 K:J
;
Figure
12.
.=
.
_.
'.á
Pet
.
li
IS
1
1
.A
., s.
;
,_ti¡i.
...,.
Tv
Clone B growing at Vancouver, B. C.
In the fall at this location it forms
a few long trailing sterile shoots.
Robust growth was noticeable.
Photographed September 2, 1964.
25
r.
°...
-
0T.
s-
N
_
----....-.-
..e:r.n
«
_2
.
..uu.Erforz *Ic
,OP
iii
-xroM,rU.1.
P
`-
yy..Y,
.
y
ra.
Figure
13.
Ligule of clone B.
i
Figure
14.
First glume of clone B.
..-
26
Glumes. Outer
-
1.7
-
2.2 mm. long,
oblong to lanceolate (Figure 14).
O. 5 - O. 6
Upper third or less of
the midnerve and edges slightly toothed.
2.
1
mm. long, 0.45
-
mm. wide;
Inner
- 1. 6 -
0.55 mm. wide; lanceolate.
Slightly toothed at tip.
Lanceolate; light grey;
Lemma.
O. 5
1. 3 - 1.
mm. wide (Figure 15).
6
mm. long, 0.4
-
Occasional unicellular
pilose hairs on nearly glabrous surface. Three nerves
reaching the apex of lemma.
Callus. Sometimes quite prominent.
Very few short hairs
forming the lateral fasicles.
Oblong -lanceolate; sometimes truncate with split apex.
Palea.
Two- nerved; 0.
6 -
0. 8
mm. long, 0.
2 -
0.
3
mm. wide.
Hyaline and partially adhering to the grain (Figure 15).
Free; enclosed in the floret; broadest at the middle;
Grain.
depressed in front. 0.9
- 1.
1
mm. long, 0.
4 -
0.
5
mm.
wide (Figure 15).
Clone
C
Source material of this clone was collected in 1955 by H.
Vaartnou from the 12th green at old Shaughnessy Heights Golf Course
at Vancouver,
B. C.
The vegetatively propagated
material has been
grown in a turf nursery in Vancouver, B. C. in the interim.
This
27
,
idc
of,¡'
a
Figure
Figure
1.5.
16.
Lemma, palea, and grain of clone B.
growing at Vancouver,
Photographed August 20, 1965.
Clone
C
B. C.
28
colonial bentgrass clone forms very fine turf and is suitable for
golf and bowling greens in southwestern British Columbia.
Under
intensive maintenance it endures moist, shady conditions with little
grain, especially when closely mowed.
Morphological characteristics
Plant. Perennial; caespitose.
Forming numerous, sparingly
branching fine rhizomes (Figure 16, 17).
25 - 60 cm. high;
Culms.
glabrous. Fertile shoots erect;
outside ones slightly geniculate at base. Occasional
stolons formed in fall by sterile shoots.
Panicle.
8 -
15
cm. long; lanceolate; brown at maturity.
Fully open at fruiting (Figure 4).
Rachis. Main nearly smooth, branches very slightly toothed.
Rachilla. Not extending beyond the floret.
Sheath closed near the base, open above.
Leaf.
usually 1.0
- 2. 5
mm. wide, 40
- 70
Blades
mm. long, nearly
linear, flat and green.
Ligule.
Slightly elongated, equal length and breadth;
long.
- 2
mm.
Truncate and toothed (Figure 18).
Glumes. Outer
- 1. 8 -
1.9 mm. long, 0.
4 - 0.
45 mm. wide.
Keeled; lanceolate; slightly toothed (Figure 19).
1. 7 -
1
1.75 mm. long, 0.4
-
0. 43 mm. wide.
Inner
-
Lanceolate;
29
Figure
17.
Pattern of rhizome and tiller growth
of clone C grown at Vancouver, B. C.
Photographed September 1, 1965.
A=I=I=1a
Figure
18.
Ligule of clone C.
30
Figure
19.
First glume
of clone C.
Figure 20. Lemma, palea, and grain of clone
C.
31
sometimes slightly toothed.
Lemma.
1. 3 -
1.42 mm. long, 0. 37 - 0.45 mm. wide;
lanceolate; silvery; nearly glabrous (Figure 20).
Three
nerves reaching the apex of lemma.
Not too prominent; slightly slanted.
Callus.
two
Palea.
lateral fasicles
0. 55 -
of
Sometimes with
hairs.
0.70 mm. long, 0.
19 -
.23 mm. wide. Oblong -
lanceolate; hyaline. Apex normally two - lipped (Figure
20).
Grain. 0.9
Partially adhering to the grain.
- 1. 0
mm. long, 0.4
- 0. 47
mm. wide.
Free,
enclosed in the floret; broadest at the middle; depressed
in front (Figure 20).
Clone
D
Source material of this clone was collected in 1955 by H.
Vaartnour from the 7th green at the old Shaughnessy Heights Golf
Course, Vancouver,
is very low growing.
B.
C., and vegetatively propagated since. It
It is
dark green in color and best adapted to
lighter, dry soil conditions. It forms an excellent, low maintenance
turf, especially in environments with great diurnal temperature
variation.
32
Morphological characteristics
Plant. Perennial; caespitose (Figure 21). Forms very few
rhizomes.
20 - 40 cm. high;
Culms.
glabrous. Fertile shoots erect,
outside ones slightly geniculate at base. Forms a few
long stolons in the fall.
Panicle.
8 -
12
Brown at maturity.
cm. long, lanceolate.
Fully open at fruiting with branchlets well separated
and spreading (Figure 4).
Rachis. Main nearly smooth, branches well toothed.
Rachilla. Not extending beyond the floret.
Sheath closed near the base, open above.
Leaf.
60
mm. long,
1. 5 - 3. 5
mm. wide.
Blades 35
-
Usually slightly
boat -shape tipped; linear and dark green.
Ligule.
O. 5 - 1. 5
Glumes. Outer
mm. long; truncate and toothed (Figure 22).
- 1.
-
7
1.95 mm. long, 0.48
Oblong - lanceolate (Figure 23).
- 0. 55
Edges and center nerve
slightly toothed one third from apex. Inner
mm. long, 0.45
-
mm. wide.
0.49 mm. wide.
- 1. 7 - 1. 85
Oblong - lanceolate
and slightly toothed.
Lemma.
1.
3
- 1. 55
mm. long, 0.
38 -
0.48 mm. wide. Oblong-
lanceolate; silvery; nearly glabrous (Figure 24).
nerves reaching the apex of lemma.
Three
3
,
-,Netsfp74"
-
G.
. 3.
I
,
*ri
V-...
..:.(1.
.
./al
U
_°,,.
t.i
.. N..'
, ',le..."
A
VI
,
I'
,.
.
,
tj,
_.; .,..1.
ti
,
.,..
0.
.-
4kkki
:1, g,
4
-
,_,
Ic
.7:
,
il...
.,,
:
ilvit
.,.
AO
,*,0
e
r
-
,...<1.:-
Figure 21. Clone D growing at Vancouver,
Photographed August 20, 1965.
.1.
-.
L
, "
.
*,,,%-
,
:"."*-1.4.1t_
--0.7!.°3` :
rit
P.0)
C
.
-
al -NI,
_.
°
;4.
Figure
22.
Ligule of clone D.
B. C.
J
34
A.
J
Figure 23. First glume of clone
D.
s
c:.
.1
,
I
Figure 24. Lemma, palea, and grain of clone
D.
35
Callus. Definitely slanted; often prominent. Fasicles of
hairs wanting.
Palea. 0.
65 - 0. 78
mm. long, 0.2
- 0. 33
mm. wide.
Oblong;
hyaline. Apex slightly two - lipped or two -lobed (Figure
24).
Grain. 0.
Partially adhering to the grain.
85 - 1.
1
mm. long, 0. 38
-
0.47 mm. wide. Free,
enclosed in the floret. Oblong; depressed in front
(Figure 24).
Clone E
Source material of this clone was collected in 1959 by H.
Vaartnou from the 4th fairway at Point Grey Golf Course, in Vancouver,
B.
C., and since vegetatively propagated in a turf nursery.
The original plant grew on heavy silty clay soil in reasonably moist
conditions.
It is light green in color and forms turf without a grain,
especially when low cutting heights are used.
Morphological characteristics
Plant. Perennial; caespitose (Figure 25). Forms new
rhizomes in favorable environment.
Culm.
40 - 60 cm. high;
glabrous.
Fertile shoots nearly
erect. Sterile shoots forming long stolons in fall.
36
1
Figure 25. Clone E growing at Vancouver,
Photographed August 20, 1965.
r-
Figure 26. First glume of clone E.
B. C.
37
Panicle.
8 - 15
Brown at maturity,
cm. long, lanceolate.
open at fruiting (Figure 4).
Rachis. Main nearly smooth, branches slightly scabrous.
Only occasionally finely toothed.
Rachilla. Not extending beyond the floret.
Sheath closed near the base, open above.
Leaf.
70
mm. long,
1. 5 - 3. 5
Blades 40
-
mm. wide, flat, nearly linear,
light green.
Ligule.
1
- 1.
5
Glumes. Outer
mm. long.
-
1.6
-
Truncate and lobed.
2.0 mm. long, 0.4
-
0.6 mm. wide.
Lanceolate; upper quarter of the center nerve and the
edges slightly toothed (Figure 26). Inner
mm. long, 0.37
-
- 1. 5 - 1. 8
Lanceolate - elliptic;
0.49 mm. wide.
slightly toothed.
Lemma.
1. 3 - 1. 6
mm. long, 0.
4 - 0. 55
lanceolate; light grey (Figure 27).
mm. wide.
Elliptic -
Three nerves reaching
the apex of lemma.
Callus. Slightly slanted; no fasicles of hairs.
Palea.
0.
6 -
0. 85 mm. long,
0.25
-
0.3 mm. wide. Lanceo-
late, tapering evenly towards two- lipped apex; loose
(Figure 27).
Grain.
1. 0 - 1.
1
mm. long, 0.
4 - 0. 55
mm. wide.
Free,
enclosed in the floret. Oblong - elliptic; depressed
in front.
38
Figure 27. Lemma, palea, and grain of clone E.
Morphological characteristics of the clones B, C, D, and E
are similar to the morphological characteristics of Agrostis tenuis
Sibth. as described by Philipson (49), except the shape of leaf tip
of clone D.
Sometimes the panicle of clone E was semiclosed at
maturity.
Specimens of all five clones are being deposited for reference
in the Herbarium at Oregon State University.
39
Field Experiment with Single Spaced Plants
Environments
Six different stations between 42o and 550 latitude were selected
to give a reasonably wide
area
to
measure the changes in morphologi-
cal features resulting from variations in environment in the five
clonally propagated genotypes.
The locations were selected to pro-
vide three daylength zones, each of which contained two locations
differing in elevation and resultant temperature conditions.
Medford, Oregon and Klamath Falls, Oregon were the southern-
most sites which had very similar day lengths but quite different
temperatures resulting from differences in elevation. Both locations
are considered to be near the southern edge of the bentgrass zone.
Vancouver, B. C. and Summerland,
B. C.
are near the 490
latitude. Vancouver was selected as a typical coastal bentgrass
growing area. Summerland has a mild, dry, sunny, interior cli-
mate, considered to be too dry for bentgrasses.
Smithers,
B. C. and
Beaverlodge, Alberta were the northern-
most sites and are normally considered to be north of the bentgrass
growing area.
These locations represent an area with cold winters,
sunny and moist summers with long days in the growing season.
Smithers represents the coastal area and Beaverlodge the northern
interior climate.
40
At the Medford experimental station the plots were established
on Meyer clay soil.
The station is situated at 42° 18' latitude;
122o 52' longitude, with an elevation of 1457 ft.
The temperature
range and rainfall totals are summarized in Table
1.
The climate
has mild winter temperatures, high summer temperatures, and
large daily temperature fluctuations.
The soil at Klamath Falls experimental station was Poe fine
sandy loam.
The latitude is 42° 12' and longitude 121° 47' having
The variation in the climate is summarized
an elevation of 4098 ft.
in Table 2.
The higher elevation
results
in lower mean
tempera-
tures than Medford. There also is less daily range in maximum and
minimum temperatures.
The experimental field of the Division of Plant Science, Uni-
versity
of B. C. was used for
Vancouver area.
establishing the field plots in the
The location has a latitude of 490 16' and a longi-
tude of 123o 15' with an elevation of 305 ft.
The soil type was an
Alderwood sandy loam, which lacked both uniformity and fertility.
Table
3
summarizes the climate. The low elevation and proximity
to the coast create a climate
characterized
by mild
temperatures
with low seasonal and daily temperature ranges.
The experimental plot at Summerland experimental
situated at an elevation of 1135 ft.
longitude of 120` 33'.
,
a
farm was
latitude of 490 34' and a
The soil type is Skaha loam,
Table
4
indicate,,
41
temperature and rainfall data at Medford
Experimental Station in the years 1964 and 1965.
Temperature
Extreme
Mean
Year
Table
1.
The monthly
and
Month
Average
°F
Low
High
Rainfall
°F
°F
in.
25
65.1
70.7
78.2
88.5
88.3
83.1
75.4
50.5
47.4
37.6
40.5
44.0
49.0
54.6
62.5
70.0
68.3
61.4
56.7
41.9
41.1
57
66
79
80
86
4.85
.33
2.22
.62
37.6
66.9
52.2
32.4
29.1
31.0
39.3
37.8
44.7
49.1
51.1
39.0
38.0
37.1
27.5
42.9
54.6
64.1
66.0
72.6
80.6
86.1
82.4
75.4
56.7
43.4
37.7
41.9
47.6
52.7
55.2
62.7
70.3
68.6
60.7
56.7
46.9
35.5
38.0
68.0
53.0
Low
High
°F
°F
30.7
25.5
31.7
32.9
38.5
46.8
51.5
48.3
39.6
38.0
33.2
34.7
44.4
55.5
56.3
1964
January
February
March
April
May
June
July
August
September
October
November
December
Mean
18
24
26
30
37
40
38
33
27
94
1.23
1.24
101
101
.74
.12
98
90
.19
.71
18
62
2.94
20
64
13.67
Total 28.86
1965
January
February
March
April
May
June
July
August
September
October
November
December
Mean
91.4
16
21
25
30
29
33
60
67
71
83
87
92
4.69
1.12
.12
3.45
.50
.75
18
28
98
95
95
31
86
24
69
1.26
.00
.40
2.17
10
53
2.85
39
39
.
Total 17.49
42
Table 2.
The monthly temperature and rainfall data at Klamath
Falls Experimental Station in the years 1964 and 1965.
Temperature
Year
and
Month
Low
Mean
High Average
°F
°F
°F
18.4
14.7
22.8
26.9
34.0
43.7
49.7
45.3
38.4
35.6
26.4
26.4
37.1
40.2
46.8
57.7
64.4
70.5
84.2
82.4
74.4
68.6
45.5
39.3
27.8
27.5
34.8
42.3
49.2
31.8
58.9
45.9
23.5
24.7
27.9
34.3
35.3
44.4
47.8
48.3
38.3
33.3
29.7
17.9
38.5
48.6
54.5
57.2
65.4
73.5
83.9
78.3
72.7
70.5
48.7
37.4
31.0
36.7
41.2
45.8
50.4
59.0
65.9
63.3
55.5
51.9
39.2
27.7
33.8
60.8
47.3
Extreme
Low
High
Rainfall
°F
°F
in.
6
13
13
55
46
68
76
24
77
32
37
34
29
90
96
91
85
24
10
82
63
7
54
1964
January
February
March
April
May
June
July
August
September
October
November
December
Mean
57.1
67.0
63.9
56.4
52.1
36.0
32.9
8
3.83
.15
.54
.25
.72
1.68
.34
.19
.12
.23
2.30
8.93
Total 19.27
1965
January
February
March
April
May
June
July
August
September
October
November
December
Mean
54
60
66
77
82
85
91
2.23
.08
.08
89
82
2.49
28
19
12
86
74
2
53
.05
2.69
1.29
5
17
18
25
24
29
38
38
1.62
.36
1.12
T
T
Total 12.01
43
Table 3.
The monthly temperature and rainfall data at Vancouver,
B. C. in the years 1964 and 1965.
Temperature
Year
and
Month
Low
Mean
High Average
°F
°F
°F
36.8
35.9
37.7
40.7
45.1
40.4
40.5
42.2
46.2
51.4
57.4
60.9
60.5
55.2
50.7
41.5
34.4
Extreme
Rainfall
Low
High
°F
°F
in.
30
50
31
32
52
9.16
3.28
64
57
74
72
78
79
67
70
52
50
4.31
1.97
2.09
2.65
2.95
1.45
5.57
2.34
6.71
6.19
1964
July
August
52.1
55.2
54.2
September
October
November
December
49.5
44.6
37.4
30.1
43.9
45.1
46.7
51.7
57.7
62.6
66.5
66.7
60.9
56.7
45.6
38.6
43.3
53.5
48.4
January
February
March
April
33.1
35.9
34.5
42.1
May
44.5
51.0
55.2
56.3
49.5
48.1
41.8
35.4
40.2
43.5
47.6
53.5
56.5
64.6
71.5
68.0
61.7
57.8
49.1
42.1
36.7
39.7
41.1
47.8
50.5
57.8
63.4
62.2
55.6
53.0
45.5
38.8
43.9
54.7
49.3
January
February
March
April
May
June
Mean
35
37
49
50
47
43
33
29
1
Total 48.67
1965
June
July
August
September
October
November
December
Mean
25
30
25
35
35
46
50
46
39
42
29
25
48
53
58
64
67
76
86
80
69
66
56
53
7.07
7.42
2.24
2.24
1.72
.67
.28
2.79
.56
7.25
5.35
7.03
Total 44.62
44
that Summerland, in the center of the Okanagan Valley, represents
a mild, dry climate having only slightly lower average
temperatures
than Vancouver and a wider range of both daily and seasonal tempera-
tures.
Smithers experimental farm is situated at 54° 44' latitude,
1270 06' longitude and its elevation is 1690 ft.
The soil type was
Telkwa clay, a very heavy clay and quite fertile.
Table
rizes the temperatures and rainfall during the seasons
5
summa-
of the ex-
periment. The lowest temperature of the experiment was experienced at this location.
Besides low summer and winter tempera-
tures, large daily temperature fluctuations are experienced.
The experimental farm at Beaverlodge has a latitude of
550 11', longitude of 119° 22' and an altitude of 2500 ft.
in the experimental
area was
This location, as Table
6
a
The soil
fertile, sandy loam, alluvium.
indicates, represents an area with cold
winter and moist summer. Both northern locations have longest
days during the summer growing season.
Clonal Increase
Single plant selections for each of the five genotypes were
made in the fall of 1963.
They were clonally propagated and grown
in the University of B. C.
,
Division of Plant Science greenhouse
during the winter of 1963 -64. Individual tillers were used for
45
Table 4.
The monthly temperature and rainfall data at Summerland
Experimental Station in the years 1964 and 1965.
Temperature
Year
and
Month
Low
Mean
High Average
Extreme
Low
High
Rainfall
in.
°F
°F
°F
°F
°F
27.8
26.8
32.4
31.9
33.4
39.3
47.3
55.4
62.8
68.6
63.7
55.8
48.8
38.0
24.0
23
43
16
20
51
65
38.9
45.5
53.0
57.8
53.6
47.2
40.7
34.0
19.6
36.0
39.9
46.2
55.7
65.3
72.5
79.4
73.7
64.3
56.9
41.9
28.3
28
35
46
64
80
50
47
39
96
84
38.1
55.0
46.6
25.7
28.6
26.6
39.4
45.9
53.5
59.6
60.0
46.7
44.1
34.3
27.0
33.2
39.1
43.3
58.8
65.9
75.7
83.7
79.3
65.4
60.8
43.3
35.0
29.5
33.9
35.0
49.1
55.9
64.6
71.7
69.7
40.9
56.9
48.9
1964
January
February
March
April
May
June
July
August
September
October
November
December
Mean
28
26
-15
82
78
70
56
44
1.03
.17
.38
.12
.21
2.25
2.33
1.02
1.37
.25
1.65
1.46
Total 12.24
1965
January
February
March
April
May
June
July
August
September
October
November
December
Mean
56.1
52.5
38.8
31.0
11
18
13
29
47
55
59
74
34
45
82
88
49
96
96
46
35
31
20
13
77
74
60
49
1.98
.29
.04
1.49
1.21
.66
.
54
2.27
.27
,
11
.86
1.45
Total 11.17
46
Table 5.
The monthly temperature and rainfall data at Smithers
Experimental Station in the years 1964 and 1965.
Temperature
Year
and
Month
Extreme
Mean
oF
Low
o
F
High
Rainfall
°F
°F
in.
18.0
-19
28.0
27.0
36.8
45.1
58.6
58.9
58.5
46.3
40.5
23.6
3.9
-
-4.8
25.7
38.2
36.7
47.8
57.4
65.5
63.8
65.0
59.7
50.4
29.9
12.5
29
45
50
60
73
80
77
82
72
73
45
42
25.6
46.1
35.9
4.5
12.3
12.6
14.6
27.0
31.1
34.9
45.3
44.3
32.7
31.2
23.0
7.6
20.0
32.6
39.5
49.3
58.7
66.9
73.8
74.3
64.1
49.2
32.6
23.4
22.6
27.1
38.2
44.9
50.9
59.6
59.3
48.4
40.2
27.8
15.5
25.7
48.7
37.2
Low
of
High
o
Average
1964
January
February
March
April
May
June
July
August
September
October
November
December
Mean
10.3
17.8
17.3
25.8
32.8
41.6
44.0
42.0
32.8
30.5
17.3
1
-14
18
21
30
33
30
21
12
-19
-42
1.33
1. 12
1.54
.
67
1.47
2.43
3. 39
2.05
1.99
1.79
2.37
1.78
Total 21.93
1965
January
February
March
April
May
June
July
August
September
October
November
December
Mean
-33
-13
-
5
15
20
23
34
29
18
21
- 4
-31
42
54
53
67
74
83
92
3.44
1.28
.
30
1.54
.93
.
50
90
74
2.06
.94
1.34
61
2. 87
46
1. 15
39
1.70
Total 18.05
47
The monthly temperature and rainfall data at Beaverlodge
Experimental Station in the years 1964 and 1965.
Temperature
Extreme
Mean
Year
High
Rainfall
High Average Low
Low
and
°F
in.
°F
°F
°F
°F
Month
Table 6.
1964
January
February
March
April
May
June
July
August
September
October
November
December
Mean
4.4
21.5
2.5
26.5
37.7
44.6
48.1
44.8
35.1
32.3
13.3
-11.6
20.6
37.0
23.0
44.4
57.3
66.3
67.7
64.4
54.6
54.8
27.5
3.4
12.5
29.3
12.8
35.5
47.5
55.5
24.9
43.4
33.7
-2.9
-4.0
8.8
27.2
35.7
43.6
49.5
50.3
34.9
33.4
7.7
5.5
12.8
18.0
29.1
43.9
58.8
66.6
71.6
71.4
51.9
52.4
21.3
18.3
5.0
7.0
19.0
35.6
47.3
55.1
60.6
60.9
43.4
42.9
14.5
11.9
24.1
43.0
33.6
57.9
54.6
44.9
43.6
20.4
-8.2
-28
0
-22
5
28
34
40
33
23
39
50
53
57
72
80
16
86
77
70
75
-22
-40
51
49
1.09
.88
.87
1.36
2.98
6.62
5.45
5.00
1.65
.65
1.65
1.01
Total 29.21
1965
January
February
March
April
May
June
July
August
September
October
November
December
Mean
-35
-26
-17
7
25
33
40
36
19
24
-22
-36
52
45
54
68
75
80
88
87
70
64
44
40
1.90
1.61
.
20
1.80
1.31
5.16
3.43
5.44
1.30
.76
1.77
.72
Total 25.40
48
increasing the number of plants.
Temperature for the days was
No
artificial light was provided.
65 °F. and
for the nights 55 °F.
Planting and Maintenance
Uniform, single tillers were planted two months previous to
the start of the experiment into standard size,
13
inch by
19
inch
flats. They were kept vigorously growing in the greenhouse at
65 °F.
day temperature and 55 °F. night temperature with no supplemental
lighting. Commercial fertilizer (10-20-10) was used every second
week. Watering was carried out as it was needed.
During 1964 the plants were space planted on three foot centers
at Medford and Klamath Falls on June 6, at Vancouver on June 24,
at Summerland on June 27, at Beaverlodge on June 30, and at
Smithe r s on July 2.
At each
location the experimental design was split plot with
main plots of observation time and subplots of clones.
plots and subplots were randomized.
Both main
Four replications were used.
Maintenance during the growing season of 1964 and 1965
summers consisted of weeding and watering.
These were carried
out by the cooperating staff of the respective experimental farms as
needed.
49
Observation Times
first morphological measurements were taken in the fall
The
of 1964.
The dates were: Summerland, September 17; Beaverlodge,
September
Smithers, September 24; Medford, October
19;
Falls, October
4;
Klamath
3;
In the spring of 1965,
and Vancouver, October 23.
the southern stations were observed before those in the north.
dates for the second measurement were: June
June
3
15
at Beaverlodge, and July
6
at Klamath Falls,
11
at Medford, June 22 at Vancouver, July
The
2
at Smithers, July
The experiment was
at Summerland.
terminated at the third observation time in the fall of 1965.
Final
data were taken at Beaverlodge on September 2, Summerland on
September
5,
Smithers on September
7,
Vancouver on September 28,
and both Medford and Klamath Falls on October 2.
Types of Observation
To get
representative leaf widths and lengths, five randomly
selected culms from each plant were collected.
fully grown, mature leaves were measured.
The two uppermost
The widest point of each
leaf and the length of the leaf from the ligule to the tip of the leaf
were measured with a micrometer to an accuracy of
ten observations per plant were averaged.
single observation per plot.
O.
1
mm.
The
This value served as a
50
The number of rhizomes per plant was counted after lifting
the plant and washing the root and rhizome clusters.
No
distinction
between the primary and secondary rhizomes was made.
If
there
were less than ten rhizomes per plant, all were measured for length
and all nodes were counted.
If the plant produced more than ten
rhizomes, then only ten were selected at random and used for observation.
The average rhizome length and the number of nodes per
rhizome were used as the observation for each plot.
To obtain the stolon number per plant all of the stolons were
counted.
For the average stolon length and the average number of
nodes per stolon, all stolons, if less than ten per plant, were meas-
ured for length and all nodes were counted. If the plant had more
than ten stolons, ten were selected at random and used for observa-
tion. The average stolon length and the average number of nodes
per stolon were used as the observation for each plot.
The
diameter of the plants was obtained by measuring the
widest and narrowest part of the plant on the soil surface, where
the tillers emerged from the soil.
The average of the two
measure-
ments was used for the average diameter of the plant.
Aerial branching when occurring at nodes above ground level
was observed.
The number of
aerial branches per plant was counted.
After lifting, the plant tops were halved at the location of both
the narrowest and the widest diameter.
Square blocks measuring
51
five centimeters by five centimeters were cut from the center of
one of the
quarter plants and all tillers within that block were
counted.
Thus density of
tillers was obtained
in unit
areas of
twenty five square centimeters.
The
facilities of the University of British Columbia Computing
Centre were used for calculating the statistical analysis of the data.
First the analysis
of variance with two
variables, having genotypes
as subplots and observation times as main plots, was calculated.
Then after regrouping the data, the second analysis of variance
was calculated using locations as the main plots and genotypes as
the subplots.
Turf Experiment
The reaction of the five clones to different mowing heights was
compared in
a
field turf trial at Vancouver, British Columbia.
The experiment was located on the Division of Plant Science
experimental field at the University of British Columbia on the
Alderwood sandy loam. All plant material required to establish
the turf plots was propagated by clonal increase in the greenhouse
starting in June 1964. The turf plots were established in September
1964 by planting the plants
regularly mowed once
cutting heights.
a
three inches apart.
week until April
1,
The turf plots were
1965 using
experimental
52
The
experimental design was split plot with four replications.
The cutting heights
represented the main plots and genotypes the
subplots. Single subplot size was twenty inches by thirty inches.
Wooden dividers of two x four lumber separated the main treatments.
They were placed to serve as a guide for the cutting height of the
lawn mower.
Five hundred grams of 10-20-10 commercial fertilizer were
applied to one hundred square feet area, two days before planting
and were worked into the upper four inches of soil.
From March
1965 to September 1965 monthly applications of five hundred
of 10-20-10 per one hundred square feet were made.
grams
As needed
all plots were irrigated at five day intervals, using one half inch
of
water per application.
In the main plots, the cutting heights
measured from the soil
surface were three eighths inch, three fourths inch, and two and
one half inches.
One uncut
area, where only the panicles were
removed when they appeared, was also included as a main plot.
The three eighths inch cutting height plots were cut every Monday
morning, Wednesday noon and Friday afternoon.
The three fourths
inch cutting height plots were cut every Monday and Friday.
The
two and one half inches cutting height plots were cut every Wednesday
throughout the growing season.
The clippings were always removed
from the plots and discarded.
The mower was a nine blade, 21 inch
53
Toro Greensmaster mower.
Morphological characteristics including tiller density, the
number of aerial branches, the weight of rhizomes and stolons per
unit area, the length of rhizomes and stolons, and the number of
rhizome and stolon nodes, were measured and recorded on observation dates of August 15, 1965, October
1,
1965 and December 1,
1965.
Sod blocks
measuring
10
centimeters square were taken in
duplicate from each subplot at each observation time and were used
for counting the aerial branches as well as obtaining the weight of
rhizomes and stolons per unit area. Ten rhizomes and ten stolons
selected at random from the individual sod blocks were used to
determine the average node length and the average node number on
both rhizomes and stolons.
Three by three centimeter square sod
blocks were used to determine the density of tillers. All sod blocks
were taken at random within the subplots.
Statistical analysis was performed at the University of British
Columbia Computing Centre.
The analysis of variance with two
variables, having genotypes as subplots and cutting height as main
plots, were calculated.
54
Growth Chamber Experiment
The five clones in this study were grown at four different night
and day temperature conditions in controlled climate conditions.
Two
Percival growth chambers, located at the Division of Plant
-
Science greenhouse at the University of British Columbia, were used
for the experiment.
Clonally propagated, uniform plants were planted one month
prior to the beginning
of the
experiment into four inch clay pots and
were kept in the greenhouse with 70oF. day and 60oF. night tempera-
tures in order to get the plants established in their respective pots.
The
cultural medium consisting of one third each of sand,
loam and Sphagnum peat moss by volume was mixed with one pound
of 10-20-10
commercial fertilizer per cubic yard before potting.
The amended soil was
sterilized and stored in
a
plastic container
until needed.
One
quarter
of an inch of washed sand was placed on the
sur-
face of the potted soil to facilitate easier watering. During the
experiment watering was done by weighing the pots twice weekly
and adding enough sterilized water to maintain the original weight.
A
split plot design with eight replications was used for this
experiment.
Main plots were temperatures and genotypes were the
subplot treatments,
55
Each replicate, consisting of five different clones in four
inch pots, was kept in a wooden frame to facilitate watering. In
this manner it was possible to lift all plants belonging to the same
replication together from the chamber, water them, and return
them to the chamber as a unit.
A 12
hour day and a
12
hour night was used in this experiment.
The illumination during daytime was 2500 f. c.
Four different temperature combinations were used:
-
45°F. at night
-
90°F. at day
2 -
60°F. at night
-
90°F. at day
3 -
45°F. at night
-
60°F. at day
4 -
60°F. at night
-
60°F. at day.
1
After growing for
60
days in the growth chambers the clones
were examined morphologically.
Characters measured included:
average plant diameter, tillers per plant, rhizome number and
length, and leaf width and length.
The analysis of variance was calculated at the University of
British Columbia Computing Centre. The analysis was calculated
with two variables having genotypes as subplots and temperatures
as main plots.
56
RESULTS
IV.
Tillering
Field Experiments with Single Spaced Plants
The number of basal
tillers (Table
7)
at Beaverlodge varied
significantly between the three observation times. The number of
tillers decreased from fall to spring. This was most pronounced
in clone A (Figure 28), which decreased from 179 to 59
25 cm.2 in this period.
Clone
C
tillers per
(Figure 29) did not suffer very
much in winter, and maintained the highest tiller number in the
The number of
spring of 1965.
1965
tillers per unit area
in the fall of
actually increased over the number of tillers in the fall of 1964.
Besides significant differences at different observation times the
tillering ability
of the five clones at this location
The descending order of the five clones ability to
varied significantly.
tiller was:
Clone C, E, D, A and B.
At Smithers (Table 7)
tillering during the first summer after
planting was slow and at the observation time of fall 1964 the clones
did not show significant differences.
Here again, clone
A
suffered
badly (Figure 30) in the winter of 1964 -65 through the activity of
Fusarium spp. The number of tillers declined from
25 cm.
2
Clones
B
136 to 44
per
and D indicated a decline in the number of basal
57
Table 7. Average number of basal tillers per 25 square centimeters
at three dates for five clones of Agrostis L. grown at six
locations.
Clone
Clone
Clone
Location
Clone
Clone
D
E
A
Average
B
C
time
Beaverlodge
Fall
179
198
282
235
258
230
59
100
239
145
139
137
177
114
558
243
284
275
138
137
359
207
227
214
136
126
184
129
137
142
44
84
247
113
155
129
206
248
386
243
347
286
129
153
272
162
213
186
190
271
474
416
442
358
91
166
285
175
194
182
204
197
571
163
268
281
162
211
443
251
301
274
83
73
159
99
101
103
Spring 1965
41
102
174
140
183
128
Fall 1965
99
135
271
134
118
152
74
103
201
124
134
128
1964
Spring 1965
Fall 1965
Average
Smithe r s
Fall 1964
Spring 1965
Fall 1965
Average
Summe rland
Fall
1964
Spring 1965
Fall 1965
Average
Vancouver
Fall
1964
Average
58
Table 7. (continued)
Location
time
Clone
Clone
Clone
Clone
Clone
A
B
C
D
E
Average
Fall 1964
263
222
412
230
189
263
Spring 1965
104
167
269
220
205
193
Fall 1965
222
162
469
291
128
254
197
184
384
247
174
237
311
255
576
346
257
349
93
156
227
201
190
174
247
205
448
276
191
273
217
205
417
275
213
265
Medford
Average
Klamath Falls
Fall 1964
Spring 1965
Fall 1965
Average
Clone Average
Fall 1964
Spring 1965
Fall 1965
Average
A
194
72
192
153
B
191
129
177
166
C
348
240
451
346
D
243
166
225
211
E
231
177
223
210
Observation average 241
157
254
217
59
1
1
Figure 28.
Clone A growing at Beaverlodge, Alberta.
This plant lacks vigor but displays typical
clone A features. Photographed September
2, 1965.
Figure 29.
Clone C growing at Beaverlodge, Alberta.
This clone is vigorous but compact.
Sterile shoots do not form stolons in fall.
Photographed September 2, 1965.
60
growing at Smithers, B. C.
This clone had not yet recovered
from previous winter damage.
Photographed July 3, 1965.
Figure 30. Clone
A
,. ,.
,
v
`
n
R'
l
v
r
,
R
e.1
r
C
.F
-I
3
Figure 31. Plants grown at Summerland, B. C.
Left to right; clones A, B, C. Photographed September 5, 1965.
61
tillers
in the spring of 1965.
an increase in
Clone E and especially clone
tillers in spring
1965 when
C
showed
compared with the fall of
Tiller number increased significantly in all of the clones in
1964.
the fall of 1965.
In addition to significant
differences at the one
percent level between the observation times and between the five
clones used, the interaction between the clones and observation
The descending
times was significant at the five percent level.
order of ability to tiller at Smithers was:
Clone C, E, D,
B
and A.
At Summerland (Table 7) all the clones;
C, D, and E;
Clones A,
particularly clones
tillered heavily in the early stages
D, and E
of the experiment,
suffered badly in the winter of 1964 -65 and the
effects of the damage were carried over until the fall of 1965
(Figure 31, 32).
The
results showed significant differences at the
one percent level between clones, observation times, and the clones
x observation times interaction.
tillers;
162
443 per 25 square
Clone
C
produced the most basal
centimeter area, and clone
A
the least;
per unit area.
At Vancouver (Table 7) the density of
tillers was
low as
compared to other locations, The differences in observation time
were not significant at the five percent level.
Significant differences
at the one percent level between the clones were recognized.
C
Clone
formed the most dense plants, having an average of 201 tillers
62
I.
.
.
L1:
Lyt
P.
«°
:.'.s
y
.
'
;i.-4.
4.4'
n
k
1
3
_
per unit area.
_
` lSgtC
ì
'R
.r.
y
p:,.
ed
Figure 32.
?..
i.t
,
i
.
^!~
-
'
.
'
,r ra
.-.
d
ii.la7t fsies,M1
Plants grown at Summerland, B. C.
Left to right; clones C, D, E. Photographed September 5, 1965.
Clone A formed the least dense plants with 74
tillers
per unit area. Interaction of clones x observation times indicated
significant differences at the five percent level. In clones
the density of
tillers decreased
D
and E
in the fall of 1965 when compared
to the spring of 1965.
Density of tillers at Klamath Falls (Table
7)
was higher than
at any other station except Summerland. Highly significant differ-
ences existed at different observation times. All the clones displayed the ability to produce more tillers per unit area in the fall
than in the spring. In addition to seasonal differences in the ability
to
tiller, significant differences between the fall
1964 and fall 1965
63
observation times were noted.
The clones displayed significant
differences in their ability to tiller. Clone
C
formed the most dense
plants with 417 tillers per 25 cm. 2, second was clone
tillers, third was clone
212
A
D
with 274
with 217 tillers, fourth was clone E with
tillers and the lowest was clone
B
with 205 tillers.
At Medford (Table 7) the clonal differences in
tillering at the
different observation times were significant only at the five percent
level. The density of tillers of all clones, except clone E, decreased
in the spring of 1965 compared with the density of
tillers
in the fall
The differences between the clones were significant at
of 1964.
the one percent level.
Especially noteworthy is the low density in
clone E in the fall of 1964 and the fall of 1965 in this environment.
Interaction between the clones and observation times was significant
at the one percent level.
tillers,
384 per unit
Clone
C
produced the highest number of
area, and clone
E had the
least,
174
tillers per
unit area.
The location totals of the five clone
tiller densities were
significantly different at the one percent level in the fall of 1964.
The highest number of
Summerland.
tillers,
358
per unit area, was produced at
This was closely followed by Klamath Falls with 349.
The intermediate group consisted of Medford with 263
Beaverlodge with 230 tillers per unit area.
tillers and Vancouver with
103
tillers per
tillers and
Smithers with
25 cm. 2
142
formed the
64
lowest group. The ability of individual clones to tiller in different
environments differed significantly at the one percent level. Clone
C
had the highest number which was 348, followed by clone
243, E with 231, A with 194, and B with 191
D
with
tillers per unit area.
Interactions (Figure 33, 34, 35) were significant at the one percent
level.
In
spring 1965 the density of tillers decreased at every location
except at Vancouver, where a small increase was recorded over the
Despite this increase, the density of tillers at Van-
fall of 1964.
couver remained lower than at the other locations. Across all
environments the five clones displayed differences.
Clone
C
formed
the most dense plants with 240 tillers, followed by clone E with 177,
D
with 166,
B
with 129, and A with only 72 tillers per 25 cm.
The capacity of the clones to
2
tiller at different environments
in the fall of 1965 was very uniform except at Vancouver which was
by far the lowest and differed significantly at the one percent level
from the others. Differences between the clones were greatest at
this observation time.
clone
C
was 451, clone
The average number of
D
192, and clone B was 177
tillers produced
was 225, clone E was 223, clone
tillers per
25 cm.
A
by
was
2
Ability to tiller, one of the most important turf characteristics,
varied widely (Table
D, 211; E, 210; B,
7)
between the clones.
166; and A, 153
Clone
tillers per
C
25 cm.
produced 346;
2
The average
65
Beaverlodge
N
500Ln
c,
400-
Fall 1964
of tillers
ci,
Fall
Spring 1965
65
1
,.. 300Number
ó 200-
0 0
o°o
ó°
óa
eao
00
0
oá
100-
z
0
,,.
C
D
O
i;
0
°
.--- ;,°
ó0
E
A
B
C
1
,P.
Ó
°Oa
A B
o
00
0 0
O
-0
D
E
A
`
0 0
0
0
C
D
B
p
p
O
°Oo
00
E
Smithers
500Ñ
0
o d
400-
\
á
-30
Fall 1964
I
Fa
Spring 1965
1
1
1b
« z00
a
z
100
A
B C
D
E
A
B
C
D
E
A
B
C
D
Clones
Figure 33. Average number of tillers of five clones
of Agrostis L. grown at Beaverlodge
and Smithers and measured at three
different observation times.
E
66
Summerland
N
U
500-
tf1
N
Ó
00
ka,400C1,
Oó
300
00
-
pp0
r-+
a
+-,
DC
48200-
0
N
.-Q
Á
100-
r
A,s;
z
°ó
600
0ó
\00o .A
Fall 1964
A
B
C
D
IA
E
0
0
Fall 1965
Spring 1965
A
B C
D
E
A B
C D E
Vancouver
500-
N
e
ó0
U
Ln
400 4
N
Fall 1964
Q300-
Fall 1965
Spring 1965
a)
+ 200w
ái
00
0
100-
o0
0°0
.0
O0
000
z
A
B
C
D
E
A
B
C
D
E
A
B
C
D E
Clones
Figure 34. Average number of tillers of five clones
of Agrostis L. grown at Summerland
and Vancouver and measured at three
different observation times.
tri
d
o
td
l
l7
Cr
w
U-1
C7
CfQ
H
op
0
0 0
0 0
00o0 060 00O 0OO0 Qo
O
o
0 0 0 C. 0
Q>° 0° 0 0 ©0 0
11111111111
°°
O
111111
cri
'v
1
opp000pop
°
rn
.0
N
O
O
IIQ'li';INIII
O
O
,
O
O
W
FP
O
O
Number of tillers (per 25 cm.
O
O
Ui
2
)
cn
H
-o
dU
Cd
0 p
o 3ßo
0
0
,
O
O
UW
0 0 0 9o0°0
ti0°000
00 y00
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0 0 0 0
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O
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O
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tA
O
CO
Un
2
cm.)
////%//////O/////////////////////
%
rn 00
C 7Q
d
O`
t U
td
u
tui
O
O
1-^'
Number of tillers (per 25
L14-euz'ejx
slip
68
number of tillers in spring was
157
compared with 241 in the fall
of 1964 and 254 in the fall of 1965.
The highest number of
tillers was produced
in the locations
normally considered outside of the best bentgrass growing areas:
Summerland, 274; Klamath Falls, 265; Medford, 237; and Beaverlodge, 214 tillers per 25 cm.
2
In
Vancouver, considered to be the
most favorable bentgrass growing area, all clones averaged only
128
tillers per
25
cm.2
Growth Chamber Experiment
The experimental clones reacted differently to different tem-
perature combinations (Table
8).
Differences in number of basal
tillers between the four temperature combinations were significant
at the one percent level (Figure 36). Clones
D
(Figure 37) and
A
(Figure 38) tillered best at 60°F. day and 45°F. night temperature.
Clones
B
(Figure 39),
C
(Figure
39), and E
(Figure 40) favored the
higher night temperature of 60°F. with 60°F. day temperature.
None of the clones
preferred the high day temperature for tillering,
except clone E. It required the higher night temperature for best
tillering and did not suffer greatly as
a
result
of a high daytime
temperature when 60°F. night temperature was used.
69
Table 8. Average number of basal tillers per plant and average
diameter of plant for five clones of Agrostis L. grown
at four different day and night temperatures.
Temperature
60°F.
60°F.
- Day
-
Night
Clone
Basal tillers
Number
A
11
B
12
C
9
41
D
9
11.
3
E
42
23
23.
7
9
68.
A
-
Day
45oF. - Night
45oF.
-
Day
Night
60°F.
-
-
Day
Night
37.4
2
36.5
7
19
29.
3
D
9
E
24
10.
18.
7
14
32.6
A
12
B
C
D
38
14
42.
44.
25.
E
24
15.
19
28.5
6
A
B
C
D
7
5
28
1
8
6
9
13.7
51.
36.
22.
3
8
6
9
2
7. 8
32
13. 4
16
26.4
A
10
B
C
D
7
32
11
58.
39.
29.
10.
E
31
17. 8
E
Average
Average
6
C
Average
90oF.
6
B
Average
60°F.
mm.
70.
39.
41.
Average
90°F.
Diameter
5
3
8
7
O
ni
A
.s;
z
d .b°
A
ww
o 0
0 if)
o, V'
o
Co
Qo
o
o0eooo00000°0°°00°0000
ooo°ooaa°
0
>to
N.
Az
s;
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O
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ww
I
N
s.zajji4 jo zaquznN
00
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00D
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nn
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0 0
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:/
0
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Illlllllillllllllllllllllllllllllllllllinllllllllllllllllll
o
llllllllllllllllllllllllllllillllll Illlllllllullllllllullll
á4
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ww
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00
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0
V'
Q)
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H
a)
ß.
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w
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cd
R
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i-i
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,
70
N
U
Z4.1
;-4
'0
a)
o
o
cd
b0
w
v`
a)
s~
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71
Figure 37. Clone D grown 60 days in the growth
chamber at 60 °F. day and 45 °F. night
temperatures.
Figure 38. Clone A grown 60 days in the growth
chamber at 60 °F. day and 45 °F. night
temperatures.
i
72
,,
r,
Sñ
-
t
,
,t-
.
Fit'
-
ta'
LA.
_
days in the growth chamber.
Top row - clone C. Bottom row - clone B.
Left column - 60 °F. day and 60 °F. night
Figure 39. Plants grown
60
temperatures.
Right column
- 90 °F.
day and 45 °F. night
temperatures.
.
=M;
-.
47.
d
Figure 40.
60 days in the growth chamber.
row
clone
D. Bottom row - clone E.
Top
Left column - 60 °F. day and 60 °F. night
Plants grown
temperatures.
Right column
- 90 °F.
temperatures.
day and 45 °F. night
73
Turf Experiment
At the August 15 observation time (Table 9) the defoliation
resulting from differences in cutting height influenced significantly
the plants' ability to produce
tillers. Three eighths inch, three
fourths inch, and two and one half inches cutting heights resulted
in more
tillers per unit area than the uncut plots. Clone
C
demon-
strated its superiority over the other clones, as it was able to
produce more tillers per unit area at all mowing heights.
average number of tillers per cm.
clone E, and
15
2
was 21 for clone C,
The
17
for
for clones A, B, and D.
At the October
1
observation time (Table
9) the
average num-
ber of tillers per unit area increased with each cutting height when
compared with uncut plots, but the differences were significant only
at the five percent level.
The differences between the experimental
clones were significant at the one percent level.
21
tillers; clone
and clone A,
13
E, 16
tillers; clone
tillers per unit area.
B,
14
The
Clone
tillers; clone
C
produced
D, 13
tiller quantity
tillers;
in all
clones decreased as compared with the first observation time.
At the December
1
observation time (Table
9)
there were
significant differences at the one percent level between the various
cutting heights.
The low cutting height (three eighths inch) reduced
significantly the number of tillers and the two and one half inches
74
Table 9. Average number of tillers per cm.2 for five clones
of Agrostis L. grown under four different cutting heights.
Observation time
Cutting height
A
13
16
11
17
7
20
21
15
15
16
16
9
E
Average
A
B
C
D
3/4 inch
E
Average
A
2
1/2 inches
uncut
16
18
13
15
23
24
18
19
19
12
18
14
14
15
21
22
14
17
16
13
18
16
1
7
8
9
9
10
19
10
10
12
A
13
11
19
13
16
14
12
12
19
13
15
14
11
15
15
21
15
13
14
9
21
18
10
B
C
D
E
Observation Average
17
17
13
16
15
13
14
22
13
15
16
11
11
18
A
10
13
19
12
14
14
8
19
Average
Average
15
14
20
E
B
C
1
13
12
18
11
13
13
Average
E
Ave rage
14
16
16
B
C
D
D
Clone
Dec.
Aug. 15
B
C
D
3/8 inch
Oct.
Clone
9
18
10
11
9
11
11
13
17
16
19
12
14
13
12
13
20
13
15
14
75
cutting height increased the number of tillers per unit area when
compared with the uncut plots. Clone
C
again produced more tillers
per unit area than any others. The average number
of
tillers pro-
duced per unit area was: clone C, 18; clone E, 11; clone D, 10; and
clones
B
and A, nine.
Diameter of Plant
Field Experiment with Single Spaced Plants
At Beaverlodge the size of the plants did not vary much (Table
10).
Clone A (Figure 28) always produced wider plants than the
other four clones. Significant differences at the one percent level
existed between the clones at all observation times.
At Smithers the plants reached their full size in the fall of
1964 (Table 10) and no significant differences between the clones
occurred at any observation time.
At Summerland the plants increased significantly in size
between the observation times (Table 10), but there were no real
differences between the clones.
At Vancouver (Table 10) there were no significant differences
between the observation times despite the large variation. Significant
differences at the one percent level existed between the clones. Clone
A
produced by far the largest plants and clone
D
the smallest ones.
76
Table 10. Average diameter of plant at three dates for five clones
of Agrostis L. grown at six locations.
Clone Average
Clone
Clone
Clone
Clone
Location
and
A
B
C
D
E
time
mm.
mm.
mm.
mm.
mm.
mm.
Fall 1964
106
61
105
109
92
95
Spring 1965
119
89
99
122
107
107
Fall 1965
189
137
145
111
141
145
138
96
116
114
114
116
101
97
65
117
125
101
Spring 1965
119
117
124
102
132
119
Fall 1965
112
102
139
102
109
113
111
106
109
107
122
111
80
55
65
105
71
75
Spring 1965
130
105
120
107
107
114
Fall
171
146
150
132
146
149
127
102
112
115
108
113
425
275
207
135
215
251
Spring 1965
577
420
315
235
192
348
Fall 1965
537
472
390
237
312
390
513
389
304
202
240
330
Beaverlodge
Average
Smithe r s
Fall
1964
Average
Summer land
Fall 1964
1965
Average
Vancouver
Fall
1964
Average
77
Table 10. (continued)
Location
Clone
Clone
Clone
Clone
Clone Average
A
B
C
D
E
mm.
mm.
mm.
mm.
mm.
Fall 1964
118
75
98
82
78
90
Spring 1965
260
127
132
110
133
152
Fall
302
160
169
167
168
193
227
121
133
120
127
145
Fall 1964
102
95
67
69
82
83
Spring 1965
212
191
140
158
154
171
Fall 1965
262
197
152
155
186
191
192
162
120
128
141
148
and
time
mm.
Medford
1965
Average
Klamath Falls
Average
Clone Average
Fall
1964
Spring 1965
Fall 1965
Average
A
156
235
264
218
B
108
163
195
155
C
101
146
193
147
D
103
139
151
131
E
110
138
168
139
Observation average 116
164
194
158
78
Continuous growth and increase in the size of plants at Klamath
Falls resulted in significant differences between observation times
at the one percent level (Table 10).
The differences between the
clones were significant at the one percent level.
Differences at the various observation times at Medford can
be easily recognized (Table 10).
All of the clones increased in size
with time. Significant differences at the one percent level between
clone A and the other four clones were observed at every observation
time.
Comparing the effect of six different environments on plant
diameter, it is noted (Table
10)
that Vancouver was the best location
for all clones at each observation time.
and clones B, D, and E to a
Clone A to a large extent,
lesser extent, favored the southern
locations over the northern ones. However, clone
C
did grow in the
north nearly as well as in the south.
Averaged over all locations and observation times, clone
was the largest and clone
D
A
was the smallest.
Growth Chamber Experiment
Four different day and night temperature combinations signifi-
cantly affected the plant size in growth chambers (Table 8). 60°F.
day temperatures were superior to the 90oF. day temperatures.
On an
average the 60°F. night temperature produced larger plants
79
than the 45oF. night temperature when the day temperature was 60oF.
But at a 90oF. day temperature, this was reversed.
Clone A pro-
duced the largest plants with only a few tillers, resulting in a very
low
tiller density when compared with clone
C
and especially with
clone E.
Growth of Leaves
Field Experiment with Single Spaced Plants
Noticeable differences in leaf width (Table 11) were observed
at Beaverlodge between the five clones, as well as between the three
observation times.
There was a definite reduction in the spring of
1965 when compared with the
results
of the fall of 1964.
All clones
showed decreased leaf width. In the fall of 1965 the leaf width of all
clones except clone
C
increased again, but remained below the width
found in the fall of 1964 (Figure 41).
At Smithers (Table 11) the
at Beaverlodge.
results were basically the same as
Significant differences between the observation
times and between the experimental clones were observed.
All the
clones had narrower leaves in the spring of 1965 than in the fall of
1964 or in the fall of 1965.
Here the leaf width of clone
C
increased
in the fall of 1965 along with the leaf width of the other clones.
the fall of 1965 the width of the leaves of clones
D
In
and E exceeded
80
Table 11. Average leaf width at three dates for five clones
of Agrostis L. grown at six locations.
Clone Average
Clone
Clone
Clone
Clone
Location
and
A
B
C
D
E
time
mm.
mm.
mm.
mm.
mm.
mm.
3.8
3.9
3.2
3.2
1.5
1.9
1.9
2.5
2.4
3.1
1.8
2.2
3.
1
2.3
2.4
2.8
2.0
2.5
2.4
3.3
2.3
3.3
1.8
2.3
4.
1
3.
3.
3.3
2.0
3.0
2.8
Beaverlodge
Fall 1964
Spring 1965
Fall
1965
Average
Smither s
Fall 1964
Spring 1965
Fall 1965
Average
2.5
2.3
3.5
3.3
1.8
1.9
2.
2.
2.3
2.4
3.5
2.9
3.7
2.9
1.8
2.5
2.4
2.7
3.4
3.5
1
3
8
1
1.9
2.4
2.5
Summer land
Fall 1964
Spring 1965
Fall
1965
Average
Vancouver
Fall 1964
Spring 1965
Fall 1965
Average
1.7
1.9
3.0
2.3
3.7
3.0
2.5
2.9
2.4
3.2
2.6
2.6
2.8
4.
2.
4.2
2.7
3.6
3.5
2.0
2.7
3.2
2.6
2.9
2.4
2.
8
3. 9
1. 8
2.7
4.6
2.4
3.6
3.9
1.4
2.7
4.2
3.
5.
2.
3.2
3.8
2.7
2.1
0
5
3.7
2.9
2.
9
3.2
3.2
3.7
3.
1
2.3
2.5
3.3
2.7
3.
1
3.4
2.5
3.0
Medford
Fall 1964
Spring 1965
Fall 1965
Average
Klamath Falls
Fall 1964
Spring 1965
Fall
1965
Average
1.5
2.4
1
2.4
3.0
3.4
5
3. 1
3. 5
4.
4.4
2.4
3.2
1.8
2.
5
1.6
4.
2.
2.8
4.0
2.4
3.3
9
1
3.
4.
1
1
8
6
2.7
3.7
2.
4.
2.
9
1
1
3.0
3.5
4.0
2.2
3.2
81
Beaverlodge
Leaf width
mm.
aiv/
_
MIMINEMI
ABCDE
mm.
ABCDE
Smithers
5-
Leaf width
ABCDE
M
=s:i
co
o
o
o
o
oc
0 O
cc
O
O
Oc
Oc
coo
0 0
0
0°0
OO
0,0
C
Fall 1964
Srin
;
ABCDE
1965
c
"V;,
ABCDE
Clones
%
-ABCDE
Fall 1965
ocs
:4 LINN c
Figure 41. Average leaf width in five clones of Agrostis
L. grown at Beaverlodge and Smithers and
measured at three different observation
times.
82
their fall
1964 widths.
The other three clones remained below the
level of the fall of 1964 (Figure 41).
At Summerland (Table 11) significant differences were observed
at the one percent level between the various observation times and
between the different clones.
The average leaf width did not decrease
in the spring of 1965, but instead increased from 2.
fall of 1964 to 2.
5
mm. in the spring of 1965 and to
fall of 1965. All clones did not perform similarly.
3
mm. in the
3.
3
mm. in the
The width of
the leaves of clones B, D, and E actually decreased in the spring
of 1965 (Figure 42).
At Vancouver (Table 11) a slight
increase in leaf width in the
spring of 1965 as compared with the fall of 1964 was observed in
every clone, except clone A. At the third observation time, all the
clones displayed a reduced leaf width (Figure 42).
Differences be-
tween individual clones were significant at the one percent level.
The average leaf width at Klamath Falls (Table 11) increased
from 3. 5 mm, in the fall of 1964 to 4.
0
mm. in the spring of 1965,
but decreased to 2.2 mm. in the fall of 1965 (Figure 43).
Differences,
significant at the one percent level, were observed at the observation
times as well as between the clones.
At Medford (Table 11) the average leaf width increased from
2.9 mm, in the fall of 1964 to 4.
then decreased to 2.
1
1
mm. in the spring of 1965, and
mm. in the fall of 1965.
Significant differences
83
Summerland
mm.
Leaf width
5-
6 O
0O0
000
600
Fa
Ç\\
A
B
C
D E
mm.
Fall 1965
Spring 1965
1964
1
C
A B
ö
0
i9
ó0
.
C
D E
OD
A B C D E
Vancouver
Leaf width
5-
00
J
00
o°o
00
coo
r,
O O
\0
DOO
NI,.
=o
A B
C
.rin
oD
D
E
A B
C
0
A
=óá%
165
ó
D
E
Fall
00
A
B
r
1965
;';'V' 0.°
C
Clones
Figure 42. Average leaf width of five clones of
Agrostis L. grown at Summerland and
Vancouver and measured at three
different observation times.
D
E
84
Klamath Falls
mm.
5-
_
0ó
Leaf width
o0
o
._\
_
f
0
0
DÓ
0e
`
o0
O
o
D
0
O
o
O
O
0
0
o
%
:% ABCDE
ABCDE
mm.
ii
i
CI
000
A
oo
ABCDE
Medford
5
Leaf width
=
4,`r
(
00
óo
co
6
00c
C
0
//
Fall\\1964
ABCDE
O O
rin 1965
O
O
O
0
0
ABCDE
ABCDE
Clone s
Figure 43. Average leaf width of five clones of
Agrostis L. grown at Klamath Falls
and Medford and measured at three
different observation times.
85
exist at the different observation times as well as between the different clones (Figure 43).
When comparing the leaf width at six locations in the fall of
percent level was observed.
1964, a significant difference at the one
perform differently in different environments
The fact that the clones
was observed (Table 11). All the clones produced their narrowest
The widest leaves for clones A and C were
leaves at Summerland.
produced at Beaverlodge; and for clones
B, D, and E
at Klamath
Falls.
In the
spring of 1965 the average station leaf widths were:
Beaverlodge, 1.9 mm. Smithers, 2.
;
0
mm.
Vancouver, 3.4 mm. Klamath Falls, 4.
;
0
,
Summerland, 2.
mm.
;
5
mm.
and Medford, 4.
;
I
These results indicate a significant increase in the leaf width
mm.
from north to south.
The leaf width (Table 11) in fall 1965 did not vary as much as
in the spring of 1965, but significant
locations.
The individual clones acted differently at different loca-
tions. Clones
the south.
differences still existed between
A and B
produced wider leaves in the north than in
The other three clones formed their widest leaves at
Summerland and Smithers, but not at Beaverlodge.
Differences significant at the one percent level in leaf length
(Table
12)
were observed at Beaverlodge at the different observation
times. In clones
A, B, and D the
leaf length was variable through
86
Table 12. Average leaf length at three dates for five clones
of Agrostis L. grown at six locations.
Clone Average
Clone
Clone
Clone
Location
Clone
and
A
time
mm.
C
D
E
mm.
mm.
mm.
mm,
B
mm.
Beaverlodge
Fall 1964
68.2
75.5
65.9
53.6
37.6
60.2
Spring 1965
52.7
70.2
65.9
52.2
53.4
59.9
Fall 1965
72.9
90.1
62.0
84.0
74.6
76.7
64.6
78.6
64.6
63.3
55.2
65.6
Fall 1964
69.1
82.9
59.7
61.2
62.0
67.0
Spring 1965
76.3
86.3
60.8
65.6
70.1
71.8
Fall 1965
48.3
63.9
83.4
58.2
93.9
69.5
64.6
77.7
68.0
61.7
75.3
69.4
81.8
110.0
73.8
71.2
93.4
86.1
Spring 1965
66.1
88.9
61.7
73.0
73.6
72.7
Fall 1965
54.7
111.8
92.6
55.6
71.0
77.2
67.6
103.5
76.1
66.6
79.3
78.7
Fall 1964
79.4
97.3
83.1
51.8
57.8
73.9
Spring 1965
92.2
98.4
63.7
53.8
44.4
70.6
Fall
71.5
82.0
96.8
65.6
72.7
77.7
81.1
92.6
81.2
57.1
57.3
74.1
Average
Smithe r s
Average
Summer land
Fall
1964
Average
Vancouver
1965
Average
87
Table 12. (continued)
Clone
Location
Clone
Clone
Clone
Clone Average
and
A
B
C
D
E
time
mm.
mm.
mm.
mm.
mm.
Fall 1964
67.9
86.2
51.5
57.9
72.3
67.1
Spring 1965
73.6
97.0
43.4
70.1
53.1
67.5
Fall 1965
62.1
79.6
64.5
61.0
68.5
67.1
67.8
87.6
53.1
63.0
64.6
67.2
67.2
93.0
57.2
64.5
76.4
71.7
Spring 1965
70.6
103.8
60.4
72.2
89.2
79.2
Fall
62.1
79.6
64.5
61.0
68.5
67.1
66.6
92.1
60.7
65.9
78.0
72.7
mm.
Medford
Average
Klamath Falls
Fall
1964
1965
Average
Average
Fall 1964
Spring 1965
Fall 1965
A
72.3
72.0
63.4
69.2
B
91.4
88.4
84.5
88.1
C
65.2
59.3
77.3
67.3
D
60.0
64.5
62.5
62.3
E
66.6
64.0
72.4
67.7
Observation average 71.1
69.6
72.0
70.9
Clone Average
88
Clone E produced the shortest leaves in the fall
the experiment.
of 1964 and the longest in the fall of 1965.
At Smithers (Table 12) clones A, B, and D produced the longest
leaves in the spring of 1965 and the shortest in the fall of 1965. The
other two clones produced their shortest leaves in the fall of 1964,
and their longest leaves in the fall of 1965.
At Summerland the longest leaves were produced in the fall of
The shortest leaves were produced there in the spring of 1965.
1964.
The different clones did not perform similarly, and differences sig-
nificant at the one percent level between the clones were recognized
(Table 12) .
No
significant differences resulted at the different observation
times at Vancouver. Differences significant at the one percent level
were observed between the clones. It is interesting to note that
clones
A
and
B
had the longest leaves in the spring of 1965 and the
other clones had the shortest leaves in the spring of 1965 (Table 12).
At Klamath
Falls clones
B, D, and E
(Table 12) produced their
longest leaves in the spring of 1965, and their shortest in the fall of
1965.
Differences, significant at the one percent level, exist at the
observation times and between the clones.
Differences, significant at the one percent level, exist between
clones at Medford.
Three of the clones;
longest leaves (Table
12) in the
A, B, and D; had
their
spring of 1965, the other two, clones
89
C
and E had their shortest leaves in the spring of 1965.
Comparing the leaf length at six different environments, dif-
ferences due to locations, significant at the one percent level, were
observed at each observation time. Besides significant differences
due to locations, differences between the clones themselves were
significant at the one percent level.
The
results (Table
11
and Table 12) indicate that the variation
in leaf width and leaf length are independent and not related to each
other.
Growth Chamber Experiment
The different clones reacted differently to the temperature
combinations used in growth chambers (Table 13). The widest leaves
were produced in 90oF. day and 45oF. night temperature combinations on clones
A, C, D, and E, but
width of clones A, B, and
D
narrowest on clone
B.
The leaf
were fairly consistent within the tempera-
ture variables of this experiment.
Leaf lengths of all clones were reduced in the 90°F. day and
60°F. night temperatures (Table 13).
Clones A, B, and
C
produced
their longest leaves with the 60°F. day and 60°F. night temperature,
and clones D and E produced their longest leaves in the 90°F. day
and 45oF. night temperature.
Clone
B
produced both the longest and widest leaf dimensions
90
Table 13. Average leaf width and leaf length for five clones of
Agrostis L. grown at four different day and night
temperatures in combinations.
Average
Average
leaf width
leaf length
mm.
mm.
Clone
Temperature
60 °F.
- Day
60 °F. - Night
A
1.7
B
C
D
2.4
E
Average
90 °F. -
Day
45 °F. - Night
A
1.7
B
C
D
2.2
1.8
1.9
E
Average
60 °F. - Day
45 °F. - Night
1.5
B
C
D
2.4
Night
52.9
77.8
40.0
Average
51.8
70.4
35.6
Average
1.5
2.2
1.2
1.7
1.3
1.6
A
1.6
B
C
D
2.3
61.5
82.5
43.4
42.8
36.5
B
C
D
E
Average
67.8
88.0
46.0
47.3
49.1
59.6
1.2
1.6
1.1
1.6
A
60 °F. -
1.8
1.9
A
E
90 °F. - Day
1.6
1.5
1.3
1.7
73.3
93.9
51.9
42.5
35.6
59.5
E
1.4
1.7
1.3
44.
8
30.1
49.
36.
1
5
31.1
45.1
91
in these
temperature combinations. Clone E produced the smallest
leaf dimensions.
Aerial Branching
Field Experiment with Single Spaced Plants
Most of the aerial branching was observed in the fall periods
(Table 14).
The
results show differences, significant at the one
percent level, between the clones, between the locations and significant interaction as well. Clones A, B, and
D
had the most aerial
branching in the Vancouver environment in the fall of 1964.
There
were great differences between the locations in the fall of 1965. At
Beaverlodge, Smithers, and Summerland all the clones produced
only occasional aerial branches. At Vancouver and Klamath Falls
all the clones except clone
D
branched freely. At Medford all the
clones branched heavily. Differences, significant at the one percent
level occurred between locations, between the clones and clones x
location interactions.
Turf Experiment
The differences in
aerial branching resulting from different
cutting heights and different clones were significant at the one percent
level (Table 15). Early in the summer the low cutting height caused
92
Table 14. Average number of aerial branches at three dates for
five clones of Agrostis L. grown at six locations.
Location
Time
Clone
Clone
Clone
Clone
Clone
A
B
C
D
E
Average
Beaverlodge
Fall 1964
3
0
0
1
1
1
Spring 1965
0
0
0
0
0
0
Fall 1965
1
0
0
5
2
2
1
0
0
2
1
1
Fall 1964
0
0
0
0
0
0
Spring 1965
0
0
0
0
0
0
Fall 1965
4
9
9
4
22
10
1
3
3
1
7
3
Fall 1964
3
1
1
3
0
2
Spring 1965
0
0
0
0
0
0
Fall
1
1
0
6
19
5
1
1
1
3
6
2
30
30
0
5
1
13
0
0
0
0
0
0
73
165
102
23
127
98
34
65
34
9
42
37
Average
Smithe r s
Average
Summerland
1965
Average
Vancouver
Fall 1964
Spring 1965
Fall
1965
Average
93
Table 14.
(continued)
Location
Time
Clone
Clone
Clone
Clone
Clone
A
B
C
D
E
Average
Medford
Fall 1964
9
1
0
0
2
2
Spring 1965
0
0
0
0
0
0
222
162
469
291
128
254
77
54
156
97
43
85
Fall 1964
1
0
0
0
0
0
Spring 1965
0
0
0
0
0
0
78
127
288
4
257
151
26
42
96
1
86
50
Fall 1965
Average
Klamath Falls
Fall 1965
Average
Clone Average
Fall 1964
Spring 1965
Fall
1965
Average
A
8
0
63
24
B
5
0
77
27
C
1
0
145
49
D
2
0
56
19
E
1
0
92
31
3
0
87
30
Observation average
94
Table 15. Average number of aerial branches per one hundred cm.2
area of five clones of Agrostis L. grown under four
different cutting heights and observed at three times.
Observation time
Average
Oct. 1
Dec. 1
Aug. 15
Clone
Cutting height
C
D
0
0
0
E
0
65
117
128
106
137
Average
0
A
0
B
C
D
0
0
A
B
3/8 inch
3/4 inch
1/2 inches
16
31
28
44
53
36
26
47
54
111
26
46
96
95
153
30
37
54
49
37
42
44
69
59
48
E
1
Average
0
116
42
53
63
62
102
23
116
143
130
184
152
95
90
134
274
78
79
117
91
116
73
177
96
115
A
6
3
35
29
39
B
C
D
76
40
45
9
E
10
49
56
77
24
34
53
52
39
46
7
60
43
37
B
C
D
E
Average
Uncut
19
129
107
A
2
0
Average
0
8
89
169
95
a
decrease in aerial branching. At the second observation time the
cutting at any height increased the number of aerial branches formed.
increased the number of
The two and one half inches cutting height
aerial branches more than the
two lower cutting heights.
At the final
observation time only the two and one half inch cutting height increased
the number of aerial branches formed over the uncut treatment.
The
three eighths inch cutting height slightly reduced the number of
branches.
Growth of Rhizomes
Field Experiment with Single Spaced Plants
Statistical analysis indicated differences, significant at the one
percent level, in the number
of
rhizomes on the different clones, in
different environments and at different seasons (Table 16). Clone
D
had an average of two rhizomes in the fall of 1964 and clone A had an
average of
31
rhizomes at the same time.
Besides the clonal differences, the environments influenced the
number of rhizomes formed. Clone
A had an
at Beaverlodge in the fall of 1964 and
87
average of six rhizomes
at Vancouver.
Clone
C
had
none at Medford in the fall of 1964, but 28 rhizomes at Vancouver
during the same period.
Each observation time influenced the number of rhizomes
96
Table 16. Average number of rhizomes per plant at three dates
for five clones of Agrostis L. grown at six locations.
Location
Time
Clone
Clone
Clone
Clone
Clone
A
B
C
D
E
Average
Beaverlodge
Fall
1964
6
1
2
0
1
2
Spring 1965
42
1
1
0
2
9
Fall
78
2
4
2
3
18
42
1
2
1
2
10
5
3
1
3
6
3
Spring 1965
19
2
4
1
1
5
Fall
86
3
104
2
6
40
36
3
36
2
4
16
7
1
3
2
2
3
21
3
41
2
4
14
175
3
93
49
5
65
68
2
46
18
4
27
Fall 1964
87
15
28
6
19
31
Spring 1965
87
23
23
1
14
30
134
34
30
1
21
44
103
24
27
2
18
35
1965
Average
Smithe r s
Fall 1964
1965
Average
Summerland
Fall 1964
Spring 1965
Fall 1965
Average
Vancouver
Fall 1965
Average
97
Table 16.
(continued)
Location
Time
Clone
Clone
Clone
Clone
A
B
C
D
Clone Average
E
Medford
Fall 1964
65
9
0
0
18
19
Spring 1965
96
4
1
1
3
21
Fall 1965
96
9
43
9
11
33
86
7
15
3
11
24
11
1
0
1
3
3
Spring 1965
227
4
7
6
2
49
Fall 1965
144
8
34
4
8
40
127
4
14
4
4
31
Average
Klamath Falls
Fall 1964
Average
Clone Average
Fall 1964
Spring 1965
Fall 1965
Average
A
31
87
118
79
B
5
5
11
7
C
6
11
52
23
D
2
2
11
5
E
8
4
9
7
10
22
40
24
Observation average
98
formed, significant at the one percent level. At Klamath Falls clone
C
formed no rhizomes in the fall of 1964, seven rhizomes in the
spring of 1965 and
34 in the
fall of 1965. At Beaverlodge, clone
formed six rhizomes in the fall of 1964,
42 in the
A
spring of 1965,
and 78 in the fall of 1965.
At every observation time clone A had the longest average
rhizomes and clone
D
had the shortest ones (Table 17).
The clonal
differences varied significantly at the one percent level at every
location and observation time. At Beaverlodge the average rhizome
length was shortest and the environment at Vancouver produced the
longest rhizomes at each observation time. Differences were sig-
nificant at the one percent level. Rhizome lengths at observation
times varied according to location and clones. Differences of locations and clones affected the length of rhizomes at Beaverlodge,
Klamath Falls, Summerland and Medford, but not at Smithers or
Vancouver.
The average number of rhizome nodes varied, and differences,
significant at the one percent level, existed between the clones and
e
nvironments (Table 18).
The interactions were significant at the
one percent level only in the fall of 1965.
largest number
The
of
rhizome nodes and clone
largest number
Clone A produced the
B
produced the fewest.
of rhizome nodes were produced at Vancouver
and the smallest number at Beaverlodge.
99
Table 17. Average length of rhizomes at three dates for five
clones of Agrostis L. grown at six locations.
Location
Clone
Clone
Clone
Clone
Clone Average
A
Time
B
D
E
C
mm.
mm.
mm.
mm.
mm.
mm.
Beaverlodge
Fall 1964
68
23
25
0
16
26
Spring 1965
84
14
27
0
27
30
118
42
64
82
35
68
90
26
39
27
26
41
Fall 1964
84
46
21
44
68
53
Spring 1965
99
48
60
44
24
55
Fall 1965
79
62
75
52
64
66
87
52
52
47
52
58
Fall 1964
79
16
41
18
45
40
Spring 1965
69
47
68
54
50
58
Fall
95
53
80
95
87
82
81
39
63
56
61
60
Fall 1964
167
84
82
46
86
93
Spring 1965
182
131
105
44
114
115
Fall 1965
117
98
97
24
100
87
155
104
95
38
100
98
Fall
1965
Average
Smithe r s
Average
Summerland
1965
Average
Vancouver
Average
100
Table 17. (continued)
Location
Clone
A
Time
mm.
Average
Clone
Clone
Clone
Clone
B
C
D
E
mm.
mm.
mm.
mm.
mm.
Medford
Fall 1964
116
37
0
0
50
41
Spring 1965
131
84
44
59
83
80
98
82
92
65
70
81
115
68
46
41
68
67
60
28
0
6
27
24
122
73
81
76
103
91
92
61
66
73
62
71
91
54
49
52
64
62
Fall
1965
Average
Klamath Falls
Fall 1964
Spring 1965
Fall 1965
Average
Clone Average
Fall
1964
Spring 1965
Fall 1965
Average
A
96
115
100
103
B
39
67
65
57
C
29
64
79
57
D
19
46
65
43
E
49
67
70
62
46
72
76
65
Observation average
101
Table 18. Average number of rhizome nodes at three dates for
five clones of Agrostis L. grown at six locations.
Location
Time
Clone
Clone
Clone
Clone
Clone
A
B
C
D
E
Average
Beaverlodge
Fall
1964
6
2
4
0
2
3
Spring 1965
7
1
1
0
3
2
Fall
7
4
6
6
3
5
7
2
4
2
2
3
9
5
2
4
7
5
Spring 1965
8
5
7
5
3
5
Fall
8
7
8
6
7
7
8
6
6
5
5
6
Fall 1964
8
2
4
3
5
4
Spring 1965
7
6
6
5
5
6
10
6
8
10
6
8
8
4
6
6
6
6
Fall 1964
10
5
6
5
6
6
Spring 1965
12
8
8
3
9
8
9
8
8
2
7
7
10
7
7
4
7
7
1965
Average
Smithe r s
Fall
1964
1965
Average
Summer land
Fall
1965
Average
Vancouver
Fall
1965
Average
102
Table 18. (continued)
Location
Time
Clone
Clone
Clone
Clone
Clone
A
B
C
D
E
Average
Medford
Fall 1964
9
4
0
0
5
3
Spring 1965
8
5
4
5
7
6
Fall 1965
9
7
8
6
6
7
9
5
4
4
6
6
Fall 1964
6
4
0
1
2
3
Spring 1965
8
6
6
6
9
7
Fall 1965
9
6
6
7
6
7
7
5
4
5
6
6
Average
Klamath Falls
Average
Clone Average
Fall 1964
Spring 1965
Fall 1965
Average
A
8
8
9
8
B
4
5
6
4
G
3
6
7
5
D
2
4
6
4
E
5
6
6
5
4
6
7
5
Observation average
103
Growth Chamber Experiment
Number of rhizomes, the rhizome length and the number of
rhizome nodes (Table
19)
varied significantly at the one percent
level between the clones. The interactions of clones to environment
were significantly different at the one percent level also. Clone
D
produced rhizomes only when the day temperature was 90°F. and
the night temperature was 45°F.
The 60°F. day and 60°F. night
temperature combination increased the number of rhizomes in clones
A and C.
Clone
A
produced the largest number of rhizomes and the
longest rhizomes in all four temperature combinations.
Turf Experiment
The weight of rhizomes per unit
area (Table 20), length of
rhizomes (Table 21) and the number of rhizome nodes per rhizome
(Table 22) varied significantly at the one percent level between clones
and observation times.
Interactions generally were not significantly
different. The amount of rhizomes produced by clone
clone E, 2.2 g.
0.
3
g.
;
clone C, 2.0 g.
;
clone B, 1.6
g.
;
A
was 9.
9
g.
;
and clone D only
All of the clones produced more rhizomes at the October
observation time than at the December or August observation times.
The uncut plots produced the
greatest mass
of
rhizomes and the
plots mowed at two and one half inches produced the least.
104
Table 19. Average number of rhizomes per plant, average
length of rhizomes and average number of rhizome
nodes per rhizome for five clones of Agrostis L.
grown at four different day and night temperatures.
Rhizome
Rhizome
nodes
length
Rhizomes
mm.
number
Number
Clone
Temperature
60°F.
-
Day
60°F.
-
Night
90oF.
45oF.
60°F.
45oF.
90oF.
60°F.
-
Day
Night
- Day
-
Night
-
Day
-
Night
Clone Average
A
11
B
C
D
2
3
0
E
1
Average
3
A
5
B
C
D
2
89.8
53.6
61.1
0
15.2
43.9
2
73.4
50.2
48.2
1
5.9
E
1
Average
2
A
3
B
C
D
2
E
0
Average
1
A
3
B
C
D
2
E
0
Average
1
1
0
1
0
A
B
6
C
2
D
0
E
0
2
6
5
5
0
1
3
4
6
4
1
20.2
39.6
2
86.9
56.2
15.0
5
0
5.3
32.7
93.7
83.4
39.4
0
13.1
45.9
85.9
60.8
40.9
1.5
13.4
3
5
1
0
1
2
7
8
3
0
1
4
6
6
4
0
1
105
Table 20. Average weight of rhizomes per 100 cm. 2 at three dates
for five clones of Agrostis L. grown under four different
cutting heights.
Weight
Observation time
Aug. 15
Cutting height
3/4 inch
2
1/2 inches
Uncut
Dec.
1
Average
g.
g.
g.
A
11.7
1.9
10.5
1.3
3.1
8.6
1.5
2.7
0.6
E
3.7
1.4
2.1
0.5
1.5
Average
1.9
A
7.5
B
C
D
2.0
0.2
E
1.5
Average
2.5
A
2.8
0.7
3.2
4.1
0.5
2.8
3.6
2.5
3.2
12.6
2.1
8.7
1.8
9.6
2.7
0.4
3.4
4.2
2.0
0.2
2.6
3.1
2.2
0.3
2.5
3.3
9.2
E
6.1
1.1
1.0
0.2
1.4
Average
2.0
2.7
6.2
0.3
1.3
0.2
1.2
1.8
7.2
0.9
1.2
0.2
1.4
2.2
A
4.
B
C
D
2.6
2.1
2.6
1.9
2.6
0.6
1.6
B
C
D
E
Average
Clone Average
1
g.
B
C
D
3/8 inch
Oct.
Clone
A
B
C
D
E
Observation Average
1.2
9
0.1
1.5
2.2
1.2
1.2
0.2
1.7
25.
1
3.2
6.7
12.
8
0.4
1.7
3.8
1.7
14.
5
2.4
2.0
0.7
2.1
4.2
5.6
1.6
14.6
1.8
9.6
1.8
0.2
1.5
2.3
0.5
2.0
0.3
2. 9
4. 4
2.1
9.9
1.6
2.0
0.3
2.2
3.
3.
2.
1
1.5
1
2
106
Table 21. Average length of rhizome at three dates for five clones
of Agrostis L. grown under four different cutting heights.
Length
Observation time
Average
Dec. 1
Oct. 1
Aug. 15
mm.
mm.
mm.
mm.
Clone
Cutting height
_
72
77
71
67
54
47
66
64
61
59
50
68
66
22
50
54
61
14
51
46
60
29
56
55
A
103
B
C
D
59
61
57
52
66
52
56
77
56
57
26
8
19
63
52
64
66
54
87
30
40
16
42
43
83
69
87
70
62
66
A
B
C
D
3/8 inch
E
Average
3/4 inch
Average
64
23
71
64
A
94
67
B
C
D
25
Average
80
66
36
72
70
A
B
73
50
C
D
69
17
67
50
55
E
66
55
E
2
1/2 inches
E
Uncut
Average
Average
61
17
59
46
10
69
50
49
74
28
42
60
B
C
D
86
66
65
32
66
50
57
19
17
E
69
60
50
50
50
A
Observation Average
64
69
54
58
45
56
23
58
53
19
59
55
74
57
60
23
60
55
107
Table 22. Average number of rhizome nodes per rhizome at three
dates for five clones of Agrostis L. grown under four
different cutting heights.
Rhizome Nodes
Observation time
Cutting height
3/8 inch
3/4 inch
Clone
Aug. 15
1/2 inches
Uncut
Average
1
Dec.
1
Average
A
8
7
8
8
5
6
7
B
C
D
8
8
8
8
6
3
2
4
E
8
6
7
7
Average
8
6
6
7
A
10
6
6
7
B
C
D
7
7
6
7
9
7
8
8
3
4
1
8
E
8
7
8
Average
7
6
6
8
6
10
6
9
8
A
2
Oct.
7
B
C
D
7
3
3
4
7
8
5
7
4
2
2
3
E
8
7
5
7
Average
7
5
5
6
A
8
6
6
7
B
5
6
8
6
C
9
7
8
8
D
2
1
3
2
E
8
7
5
7
Average
6
5
6
6
A
9
6
7
7
B
C
D
7
6
6
6
8
7
7
7
4
2
2
3
E
8
7
6
7
7
6
6
6
Observation Average
108
Clone A produced the longest rhizomes, 74 mm.
by clones C and E with 60 mm.
;
B, 57 mm.
;
,
followed
and D, 23 mm.
Cutting
height had no influence on rhizome length.
At individual observation times the number of rhizome nodes
varied significantly by clone differences. Cutting height had no influence on number of rhizome nodes.
Clone D produced fewer rhi-
zome nodes than the other clones.
Growth of Stolons
Field Experiment with Single Spaced Plants
In this
experiment none of the clones formed stolons in the fall
of 1964 and the spring of 1965.
In the
fall of 1965 differences signifi-
cant at the one percent level occurred between the clones (Table 23)
None of the clones formed stolons at Med-
and between locations.
ford. At Beaverlodge and Klamath Falls only occasional stolons
developed. At Vancouver all of the clones had stolons, especially
clones
A, D, and E.
At Smithers clones A, B, and D formed oc-
casional stolons while clones
C
and E did not.
occurred at Summerland where clone
clone D, 153, and clone
C
A
The greatest variation
formed an average of 306,
no stolons per plant.
Differences in the length of the stolons (Table 24) varied with
clones and locations.
The longest stolons formed were an average
109
Table 23. Average number of stolons at three dates for five
clones of Agrostis L. grown at six locations.
Location
Time
Clone
Clone
Clone
Clone
Clone
A
B
C
D
E
Average
Beaverlodge
Fall
1964
0
0
0
0
0
0
Spring 1965
0
0
0
0
0
0
Fall
4
1
1
4
1
2
1
0
0
1
0
1
0
0
0
0
0
0
0
0
1965
Average
Smithe r s
Fall
19 64
Spring 1965
0
0
0
49
16
5
13
0
2
0
0
4
0
13
4
0
0
0
0
0
0
0
0
0
0
0
0
306
6
0
153
1
93
102
2
0
51
0
31
Fall 1964
0
0
0
0
0
0
Spring 1965
0
0
0
0
0
0
16
Fall
1965
Average
Summe r land
Fall 1964
Spring 1965
Fall 1965
Average
Vancouver
0
Fall 1965
55
3
3
8
Average
Medford
18
1
1
3
13
4
0
0
0
0
0
0
Spring 1965
0
0
0
0
0
0
Fall
19 65
0
0
0
0
0
0
Average
Klamath Falls
0
0
0
0
0
0
Fall 1964
0
0
0
0
0
0
Spring 1965
0
0
0
0
0
0
Fall 1965
Average
2
0
1
4
0
1
1
0
0
1
0
1
Fall
19 64
5
110
Table 24. Average length of stolons at three dates for five clones
of Agrostis L. grown at six locations.
Clone
Clone
Clone
Clone Average
Location
Clone
D
E
A
B
C
Time
mm.
mm.
mm.
mm.
mm.
mm.
Beaverlodge
Fall 1964
0
0
0
0
0
0
Spring 1965
0
0
0
0
0
0
282
131
75
178
113
156
94
44
25
59
38
52
0
0
0
0
0
0
0
0
0
0
0
0
203
223
0
185
0
122
68
74
0
62
0
41
Fall 1964
0
0
0
0
0
0
Spring 1965
0
0
0
0
0
0
389
379
0
278
136
236
130
126
0
93
45
79
Fall 1964
0
0
0
0
0
0
Spring 1965
0
0
0
0
0
0
237
145
146
108
128
153
79
48
49
36
43
51
Fall 1965
Average
Smithe r s
Fall
1964
Spring 1965
Fall
1965
Average
Summerland
Fall 1965
Average
Vancouver
Fall 1965
Average
111
Table 24. (continued)
Clone
Location
A
Time
mm.
Clone
Clone
Clone
Clone
B
C
D
E
mm.
mm.
mm.
mm.
Average
mm.
Medford
Fall 1964
0
0
0
0
0
0
Spring 1965
0
0
0
0
0
0
Fall 1965
0
0
0
0
0
0
0
0
0
0
0
0
Fall 1964
0
0
0
0
0
0
Spring 1965
0
0
0
0
0
0
175
0
59
104
0
68
58
0
20
34
0
23
Average
Klamath Falls
Fall 1965
Average
Clone Average
Fall 1964
Spring 1965
Fall 1965
Average
A
0
0
214
71
B
0
0
146
49
C
0
0
47
16
D
0
0
142
47
E
0
0
63
21
112
of 79 mm. per stolon at Summerland, while none were produced at
Medford.
Clone A produced the longest, averaging
shortest stolons, averaging
The
21 nana.,
71
mm.
were formed on clone
The
C.
differences between clones and between location, displayed
significance at the one percent level.
The number of stolon nodes (Table 25) varied significantly at
the one percent level between clones and between locations.
Clone
A
produced an average of three nodes per stolon while clones
E
produced an average of only one node per stolon. More stolon
C
and
nodes per stolon were produced at Beaverlodge than at other loca-
tions.
Turf Experiment
At the
no
first observation time (Table
26) in the summer of 1965,
significant differences, at the five percent level, existed.
fall, at the October
1
observation time, clone
D
In
early
produced more stolon
mass than the other four clones. It also had the largest mass during
the third observation.
The two and one half inches cutting height
increased the amount of stolons in clones
produced more stolons for clones
A
B
and E,
and D, while uncut plots
Differences between the
clones were significant at the one percent level with clone
ing the
largest stolon mass.
D
produc-
113
Table 25. Average number of stolon nodes at three dates for
five clones of Agrostis L. grown at six locations.
Location
Time
Clone
Clone
Clone
Clone
Clone
A
B
C
D
E
Average
Beaverlodge
Fall 1964
0
0
0
0
0
0
Spring 1965
0
0
0
0
0
0
11
6
6
7
8
8
4
2
2
2
3
3
Fall 1964
0
0
0
0
0
0
Spring 1965
0
0
0
0
0
0
11
10
0
11
0
6
4
3
0
4
0
2
Fall 1964
0
0
0
0
0
0
Spring 1965
0
0
0
0
0
0
12
9
0
11
3
7
4
3
0
4
1
2
0
0
0
0
0
0
Spring 1965
0
0
0
0
0
0
Fall 1965
6
4
3
4
6
4
2
1
1
1
2
1
Fall 1965
Average
Smithe r s
Fall
1965
Average
Summerland
Fall 1965
Average
Vancouver
Fall
1964
Average
114
Table 25. (continued)
Location
Time
Clone
Clone
Clone
Clone
Clone
A
B
C
D
E
Average
Medford
Fall 1964
0
0
0
0
0
0
Spring 1965
0
0
0
0
0
0
Fall 1965
0
0
0
0
0
0
0
0
0
0
0
0
Fall 1964
0
0
0
0
0
0
Spring 1965
0
0
0
0
0
0
11
0
9
9
0
6
4
0
3
3
0
2
Average
Klamath Falls
Fall 1965
Average
Clone Average
Fall 1964
Spring 1965
Fall
1965
Average
A
0
0
9
3
B
0
0
5
2
C
0
0
3
1
D
0
0
7
2
E
0
0
3
1
0
0
5
2
Observation average
115
Table 26. Average weight of stolons per 100 cm. 2 area at three
dates for five clones of Agrostis L. grown under four
different cutting heights.
Observation time
Average
Dec. 1
Oct. 1
Aug. 15
height
Cutting
Clone
g.
g.
g.
g.
3/8 inch
3/4 inch
2
1/2 inche s
Uncut
Clone Average
0.25
1.50
0.25
2.31
0.62
0.99
0. 12
0. 16
0.50
0.19
2.81
0.81
0.89
0.71
0.15
1.75
0.48
0.65
0
Average
0.06
0.04
0.31
1.31
0.44
3.31
0.62
1.20
1.50
0.31
4.62
0.69
1.42
0.14
0.94
0.25
2.64
0.46
0.79
A
0. 12
0. 25
0. 12
0. 16
B
C
D
0.56
1.25
0.44
8.75
E
0
Average
0.15
4.81
0.25
8.62
0.62
2.91
49
2.21
0.23
5.79
0.83
1.85
A
0.06
B
C
D
0
2.50
0.56
2.06
3.19
1.54
1.25
0. 18
0. 31
3. 06
1. 18
0.06
3.44
E
0.
Average
0.06
1.12
1.59
12.69
4.12
5.02
5.39
1.75
2.22
A
0. 1 1
0. 17
0.83
0.58
2. 05
1. 61
0.05
0.06
0.01
0.31
4.42
1.00
7.22
1.87
0.51
1.28
0.45
3.90
0.88
A
0. 12
B
C
D
0.12
E
0
Average
0.07
A
0.12
B
C
D
0
E
B
C
D
E
0
0. 12
0
0
0
0.06
0.75
1.87
2.
116
Cutting height and clones significantly influenced the stolon
length only at the last two observation dates (Table 27). Clone
D
produced the longest stolons and the uncut plots produced longer
stolons than other mowing heights.
Cutting height signficantly influenced the number of stolon nodes
only at the December observation time (Table 28).
The uncut plots
had a much larger number of nodes than the mowed plots.
Clonal
differences were significant during the latter two observation times.
Clone
least.
D
produced the most stolon nodes, while clone
A
produced the
117
Table 27. Average length of stolon at three dates for five clones
of Agrostis L. grown under four different cutting heights.
Observation time
Aug. 15
Cutting height
3/8 inch
3/4 inch
1/2 inches
Clone Average
Dec. 1
mm.
Average
mm.
12
16
35
29
16
11
55
42
32
42
22
24
0
12
30
29
25
20
43
27
26
A
B
21
14
C
D
0
11
15
37
18
59
E
0
24
Average
9
31
A
8
B
C
D
5
7
0
27
49
28
7
Average
6
69
39
42
60
E
10
Clone
B
C
D
E
Uncut
1
mm.
A
2
Oct.
mm.
35
30
0
67
27
22
60
41
23
97
90
0
52
53
34
30
Average
17
33
47
A
66
204
246
B
C
D
0
82
22
E
0
157
221
188
139
190
Average
18
92
162
67
121
A
B
C
D
32
64
12
7
59
41
70
70
76
9
97
98
2
41
67
E
3
22
52
23
70
28
39
172
79
112
118
69
110
55
47
41
68
37
118
Table 28. Average number of stolon nodes per stolon at three dates
for five clones of Agrostis L. grown under four different
cutting heights.
Observation time
Cutting height
3/8 inch
3/4 inch
Clone
Aug. 15
A
0.9
1.4
B
C
D
1/2 inche s
Uncut
Average
0.9
E
0
Average
0.6
A
0.5
0.5
0.5
B
C
D
0
Average
0.7
0.4
A
B
1.2
0.9
C
D
0
E
0
Average
0.6
E
2
0
0.7
Oct.
1
Dec.
1
Average
1.0
1.2
3.6
2.4
5.8
3.2
3.2
3.6
2.4
1.0
2.9
1.6
6.2
5.7
3.8
4.3
3.0
2.5
3.0
4.1
3.3
7.1
3.8
4.2
0
1.2
3.3
3.5
6.6
5.3
3.7
2.7
2.4
4.6
3.3
2.8
0.5
1.2
1.0
3.3
1.3
6.6
3.1
3.0
3.3
3.4
2.5
5.1
3.2
8.7
6.9
8.4
10.7
5.0
3.4
4.1
6.
9
5.7
4.
1
1.6
4.7
2.9
2.6
A
1.2
B
C
D
0
E
0
8.1
3.2
Average
0.7
4.4
9.3
8.8
4.2
4.5
A
1.0
B
C
D
0.7
0.5
0.6
0.2
2.4
3.6
2.4
2.8
4.3
4.4
2.1
2.6
6.9
3.3
7.6
6.5
E
1.5
0.7
2.5
6.
5
2.4
5.0
3.3
119
DISCUSSION
Five genotypes of Agrostis L. were used in this study to
determine the response to variations in environment. At the beginning, due to a lack of comprehensive studies in the field of Agrostis
sp.
,
all of them were classified as belonging to the species Agrostis
tenuis Sibth. After doing
tis
L.
,
a
taxonomic study in 1965 on genus Agros-
the reliability of the botanical names used in reference litera-
ture for genotypes employed in this study were doubted, and soon it
was clear that clone A did not belong to the species Agrostis tenuis
Sibth.
Its macromorphological characteristics are similar to the
species Agrostis gigantea Roth. as described by Philipson (49). The
morphological characteristics of the other four clones are quite
similar to those
(49).
of
Agrostis tenuis Sibth.
as described by Philipson
Detailed taxonomic study of the populations from which the
clones were selected is warranted.
Tillering
In this study the density of
tillers varied significantly between
the clones in every location at each observation time.
This result
agrees with Mitchell (41) in that it is an hereditary characteristic
and varies according to the species or clone.
characteristic and depending
Being an hereditary
on the number of bud
primordia formed,
120
its ultimate expression is influenced by environment. Significant
differences between locations indicate that the final density of tillers
would be decided simultaneously and collectively by all factors of
the environment at the time that the bud primordia are formed.
The
(35).
decrease in tiller number in the summer agrees with Langer
In
this study the decrease was caused mainly by the fact that
some of the tillers developed in the previous fall or early spring
completed their life cycle, and secondly by death due to winter
diseases.
ous
In
spring when fertile culms developed on healthy vigor-
tillers, the weak tillers died and there were
in late spring.
It is possible that high
few new ones formed
temperatures
in spring and
summer inhibited the development of lateral buds (41).
The remark-
able increase in the density of tillers in the fall can be accounted for
by the effect of short days in combination with low night
(44).
The
temperatures
overall low density of tillers at Vancouver can be ex-
plained by comparatively low soil fertility at that location (21, 35,
53, 67)
especially by
a
possible shortage of phosphorus (9, 22) and
nitrogen (12, 30). Another factor which might have influenced tiller ing was the lower light intensity at the time of development of bud
primordia (40,
The
53, 41).
results
of this study did not
agree completely with the
Nittler, Kenny and Osborne (46) observations that variety differences
are small when light is available in sufficient quantities and large
121
when light is the limiting factor.
In
this study the variety differ-
ences were greatest at Summerland, Medford and Klamath Falls
In field conditions the
and least at Vancouver (Table 7).
variety
differences of certain plants in tiller density can be more clearly
demonstrated in the more sunny locations.
The growth chamber experiment demonstrated that different
clones of Agrostis sp. reacted differently to environments provided
in this experiment (Figure 40).
Clones
A
and D formed most
per plant in 60°F. day and 45°F. night temperature.
tillers
The other
clones tillered most frequently in the 60°F. day and 60°F. night
temperature. Clones
E and D had
completely different growth habits
in growth chambers (Figure 40) when compared with field experi-
ments, demonstrating the need to use extreme caution when applying
results obtained under artificial conditions to field research.
In the
turf experiment the three fourths inch and two and one
half inches cutting heights increased the density of tillers at each
observation time (Table 9).
In
three eights inch cutting heights
there were significant differences between the clones. Low cutting
height reduced the density of tillers of clone A at each observation
time. However, the density of tillers of the other four clones was
increased by low cutting heights in the summer and early fall but
having its tiller density reduced
decreased in late fall. With clone
A
by low cutting height, it cannot be
regarded as
a clone for
turf
122
requiring low, frequent mowing. The other four clones would be
valued as better varieties for turf requiring intensive maintenance.
Low cutting height in late fall reduced the quality of
turf in every
clone.
The plant size varied significantly between both locations and
between clones. Clone
formed larger plants than the other clones.
A
This can be related to its ability to form extended rhizomes which
emerge readily and tiller only slightly, thus forming a plant with
low
tiller density and large diameter.
It favors southern locations,
because in northern areas Fusarium sp. attacks this clone in both
fall and spring causing severe damage (Figure 44).
Clones B, D,
and E reacted most favorably to the southern locations.
Clone C,
being the most vigorously tillering clone, reacted better than any
other clone to widely diversified environments.
It was the best
performing clone at every location, particularly at Beaverlodge
(Figure 29) and Summerland (Figure 45).
tiller density indicate that clone
than the other four clones.
C
Clonal differences in
is far better for turf purposes
Clone A produces very weak turf at
springtime in areas where Fusarium sp. causes severe damage to
it.
Growth of Leaves
The leaf width varied significantly between locations and clones
123
in the fall of
1964
(Table 11).
The final leaf width depended upon
the environment and upon the genetic structure of the plant.
In the spring of 1965 (Table 11) the average width of the leaves
of all clones displayed a definite trend,
at Beaverlodge to
4. 0
mm. in Medford.
increasing from 1.9 mm.
At Summerland, Vancouver,
Klamath Falls and Medford (Figure 42, 43) the average width of
leaves in the spring exceeded the width of leaves in the fall; a fact
which can be explained by increased light intensity and day length (19 ).
Contrary to this, at Beaverlodge and at Smithers (Figure 41), the
average leaf width in the spring was far below the width of the leaves
in the fall.
The reduction in leaf width at the two northern stations
may be explained not only by increased temperatures, but also by
the long, cold vernalization effect at the time of development of the
leaf primordia.
In the growth
chamber experiment, significant differences in
leaf width existed between the clones.
The clone x temperature
interactions were also significantly different. Clone
narrowest leaves in
90 F. day
temperature (Table
B
13).
produced the
The other
four clones produced widest leaves in the 90oF. day and 45oF. night
temperature. Narrowest leaves were produced in the 60°F. day
and 45oF. night temperature.
a
This indicates that clone
lower day temperature to produce widest leaves.
B
requires
In addition to
high day temperatures, the other four clones require a low night
124
temperature for widest leaf growth.
The length of leaves depends mainly upon the genotype of the
clone. In addition to the genetic control, environment was an im-
portant factor in modifying the leaf length. Differences existed at
different locations. The fact that the clones performed differently
at various locations confirms the suggestion (11) that leaf length is
governed by multiple pairs of genes. Increases in day length in-
creased the length of leaves. This, along with mineral nutrition,
was the most significant environmental factor affecting length of
leaf growth. Reduction of the leaf length at Summerland, Vancouver,
and Medford in the spring of 1965 was related to the increase in light
intensity which especially affected clones
C
and E.
No
clear relation-
ship between the leaf width and leaf length was observed.
Aerial Branching
Significant differences occurred at various locations, indicating that aerial branching is controlled mainly by environment.
Aerial branching occurred in the fall (Table
14)
mainly in environ-
ments where the fertile shoots did not die back to the ground level
in the fall.
The branching occurred by nodes above the soil surface,
as soon as the apical bud dominance was diminished.
Then the
highest lateral buds started to grow, either as aerial branching
or if at ground level or below, as sterile shoots.
125
In the
turf experiment (Table
to be a definite factor in influencing
15) the
cutting height was found
aerial branching. The two and
one half inches cutting height increased the
aerial branching signifi-
cantly at each observation time with all clones in this experiment.
The three eights inch and three quarters inch cutting heights (Table
15)
also increased aerial branching in early fall, but not in late fall.
This indicates that short daylengths (20) would not be the main reason
for the activity of the lateral buds.
of
Clone
C
had the highest number
aerial branches per plant. The relationship between the clones
used in this experiment regarding ability to form aerial branches
was the same as the ability to tiller, indicating a possible associa-
tion between those two characteristics.
Growth of Rhizomes
The genotypes varied in
their ability to form rhizomes (Table
16).
Clone A had more rhizomes than any other clone.
clone
C
could be classified as plants with rhizomes.
It and
The other
three clones would be classified as plants with occasional rhizomes.
Besides clone differences the environment seems to be important in
determining the number of rhizomes formed. Significant differences
between locations can be pointed out, especially with clone
The number of rhizomes
cially in clones
A
and C.
A
and C.
increased with the age of the plant, espeShort daylength in the fall increased the
126
number of rhizomes, especially at northern stations, but not in
clone A at Medford and Klamath Falls.
In the growth chamber experiment, clone A produced more
This relates directly to the field
rhizomes than the other clones.
experiment results where clone
A
also produced most rhizomes.
Clonal difference in the turf experiment was the significant
factor affecting rhizome production.
A
change in cutting height did
not change the number of rhizomes in summer, but defoliation in
the fall reduced the amount of rhizomes formed.
Significant differences between the clones and between the
locations used in this study indicate that length of rhizomes has
a
genetic control, but that the environment is a very important factor
also affecting it. Significant interaction between the clones and
environments were demonstrated. Long daylengths in the spring
increased the length of rhizomes in clone
four clones.
A, but not in the
other
It is possible that the short daylength at Beaverlodge
in the fall of 1965 combined with other environmental factors, such
as mineral nutrition, reduced the length of rhizomes of clones B,
C, D, and E
at Beaverlodge.
Temperature, in growth chambers, and cutting heights, in the
turf experiment, did not affect the length of rhizomes significantly.
In both
experiments, only significant differences existed between the
clones used.
In every experiment in this study the number of
127
rhizome nodes varied according to the length of rhizome.
No
correlation between the plants ability to tiller and to form
rhizomes, despite both of them being lateral meristems, was observed.
Growth of Stolons
This study agrees with Ryle and Langer (51) in that the formation of stolons can be controlled by photoperiods.
The stolons formed
only in the fall when short days with low light intensity occurred. In
addition to environmental control (Table 23) the clones, having he-
reditary differences, in times of limited reproductive and accentuated
vegetative growth, can produce stolons or tillers. Significant differences between the different clones existed in addition to the significant variations between the various environments.
Clones A and
B, E, and
D
especially
were able to form more stolons than clones
C.
No
association between the ability to tiller
and form stolons were observed.
Negative association between the
clones ability to form turf without grain and to produce stolons can be
pointed out.
128
SUMMARY
The genus Agrostis L.
,
which is one of the most important
genera for turf production in Northwest America, appears not to
have been given intensive study in this area.
The morphological
features, such as tillering, diameter of plant, growth of leaves,
aerial branching, growth
of
of
rhizomes and stolons of five genotypes
bentgrass, were compared under various environmental condi-
tions in this work.
Clonally propagated material of five greatly different genotypes of bentgrass were used in six environments located from
southern Oregon to northern British Columbia.
The same clones
were studied under different temperatures in growth chambers, as
well as under different cutting heights in turf plots.
The following
conclusions can be drawn from the results obtained in this study:
Morphological characteristics of bentgrasses varied in their
responses both to genotype and environment.
The ability to
genotypes.
tiller
in
bentgrasses varied widely between
Disease susceptibility is one of the factors important
to the density of
tillers, especially
in turf.
Fusarium sp. diseases
reduced considerably the density of tillers in bentgrasses.
The density of
tillers depended
on the season.
Normally in the
summer the density of tillers was reduced because of limited initiation of bud primordia in early summer.
Temperature, especially
129
night temperature, had a definite effect on the density of tillers.
The genotypes
varied widely in their requirements for optimum
temperature for tillering. Defoliation affected the density
of
tillers.
The optimum cutting height depended upon genotypes and environ-
ments. Short days in the fall increased the density of tillers.
All the clones tillered more profusely at locations with higher
light intensity.
The diameter of the plants depended upon both the genotypes and
the environments.
Environments with ample rainfall in fall and
spring, when the plants' vegetative growth was vigorous, produced
low
tiller density and larger diameter plants. The different species
and varieties can perform differently under similar environments.
Increased light intensity and day length were factors in increasing the leaf width at locations where the temperature had not affected
the development of leaves or the initiation of leaf primordia.
Tem-
perature combinations had different effect on genotypes. The same
temperature combination increased the width of leaves in one genotype and decreased it in others.
The length of leaves depended mainly upon the genotype.
No
correlation between the leaf width and length was observed.
Aerial branching occurs when the dominance of the apical bud
is removed, and when the
the tillering and
lateral buds develop. Correlation between
aerial branching
in clones was
observed.
130
No
correlation existed between the clones'ability to form
tillers and rhizomes despite the fact that both
of them are
classified
as lateral meristems.
Short days and decreased light intensity in the fall combined
with low night temperatures are proposed to be the main environ-
mental factors which accentuate the formation of stolons,
The
ability to form stolons was not correlated with the plant's ability to
form rhizomes or tillers.
131
BIBLIOGRAPHY
The effects of cutting, light intensity and night
temperature on growth and soluble carbohydrate content of
Lolium perenne L. Plant and Soil 8:199 -230. 1957.
1.
Alberda, Th.
2
Allard, H. A. and M. W. Evans. Growth and flowering of some
tame and wild grasses in response to different photoperiods.
Journal of Agricultural Research 62:193 -228. 1941.
3.
Ashby, E,
Phytologist
Studies in the morphogenesis of leaves. New
47 (2):153 -176. 1948.
4.
Bean, E. W. The influence of light intensity upon the growth
of an S. 37 Cocksfoot (Dactylis glomera.ta) Sward, Annals of
Botany 28:427-443. 1964.
5,
Beard, James B. Factors in the adaptation of turfgrasses to
shade. Agronomy Journal 57:457 -459. 1965.
6.
Beard, James B. and William H. Daniel. Effect of temperature
and cutting on the growth of Creeping Bentgrass (Agrostis palus tris Huds) roots. Agronomy Journal 57:249 -250. 1965.
7.
Beinhart, George. Effects of environment on meristematic
development, leaf area, and growth of white clover, Crop
Science
8.
3
(3):209 -213.
Bjorkman,
9.
O.
S.
40:254 -258.
1963.
Chromosome studies in Agrostis II.
Hereditas
1954.
Brenchley, W. E. The phosphate requirement of barley at
different periods of growth. Annals of Botany 43:89-100. 1929.
10.
Effects of drought, temperature, and nitrogen
on turfgrasses. Plant Physiology 18:19 -36. 1943.
11.
Clausen, J. C. D. D. Keck and W. M. Hiesey. The effects of
varied environments on plants. Washington, D. C. 1940. 452
p. (Carnegie Institution of Washington. Publication 520)
Carrol, J.
C.
,
,
12.
Studies on growth and development in Lolium.
II. Pattern of seed development of the shoot apex and its ecological significance. Journal of Ecology 39:228 -270. 1951.
Cooper, J. P.
132
13.
Cowett, Everett R. and Milton A. Sprague. Factors affecting
tillering in alfalfa. Agronomy Journal 54(4):294 -297. 1962.
14.
Darrow, R. A. Effects of soil temperature, pH, and nitrogen
nutrition on the development of Poa pratensis. Botanical Gazette 101:109 -127. 1939.
15.
Davis, Larry A. and Horton M. Laude. The development of
tillers in Bromus mollis L. Crop Science 4(5):477 -480. 1964.
16.
Evans, Morgan W.
1927.
55 p.
(U. S.
history of timothy. Washington,
Department of Agriculture. Bulletin 1450)
The life
17.
Evans, M. W. and J. E. Ely. Growth habits of Reed Canary
grass. Agronomy Journal 33:1017-1027. 1941.
18.
Evans, M.
The growth of Kentucky
in
late spring and in autumn
bluegrass and of Canada bluegrass
as affected by length of day. Journal of the American Society
of Agronomy 31:767-774. 19 39
W. and J. M.
Watkins.
.
19.
Friend, D. J. C. , V. A. Helson and J. E. Fisher. Leaf growth
in Marquis wheat as regulated by temperature, light intensity,
and daylength. Canadian Journal of Botany 40(10):1299 -1311.
1962.
20.
Garner, W. W. and H. A. Allard. Further studies in photo periodism. The response of the plant to relative length of day
and night. Journal of Agricultural Research 23:871 -920. 1923.
21.
Gericke, W. F. Some effects of successive cropping of barley.
Journal of the American Society of Agronomy 9:325 -332. 1917.
22.
Grantham, A. E. Tillering as a factor in determining the
desirable qualities of winter wheat. Proceedings of the American Society of Agronomy 4:75 -81. 1912.
23. Hansel, H. Vernalisation of winter rye by negative temperatures and the influence of vernalisation upon the lamina length
of the first and second leaf in winter rye, spring barley, and
winter barley. Annals of Botany 17:417 -432. 1953.
24.
Hanson, A. A. and F. V. Juska. Winter root activity in
Kentucky bluegrass (Poa pratensis L. ) Agronomy Journal 53:
372 -374. 1961.
133
25.
Harrison, C. M. Responses of Kentucky bluegrass to variation
in temperature, light, cutting and fertilizing. Plant Physiology
9:83 -106.
1934.
26.
Hiesey, W. M. Growth and development of species and hybrids
of Poa under controlled temperatures. American Journal of
Botany 40:205 -221. 1953.
27.
Hiesey,
28.
Jowett, D. Population studies on lead -tolerant Agrostis tenuis.
Evolution 18(1) :70 -81. 1964.
29.
Juska, Felix V. Shade tolerance of bentgrasses.
Reporter 3:28 -34. 1963.
30.
Kershaw, K. A. An investigation of the structure of a grassland
community. I. The pattern of Agrostis tenuis. Journal of
Ecology 46:571 -592. 1958.
31.
Kershaw, K. A. An investigation of the structure of a grassland
community. II. The pattern of Dactylis glomerata, Lolium
perenne and Trifolium repens. III. Discussion and conclusion.
Journal of Ecology 47:31-53. 19 59
and H. W. Milner. Physiology of ecological
races and species. Annual Review of Plant Physiology 16:
203 -216. 1965.
W. M.
Golf Course
.
32.
Knight, R. The relation between tillering and dry matter production in Cocksfoot (Dactylis glomerata L. grown under
spaced and sward conditions. Australian Journal of Agricultural Research 12:566 -577. 1961.
)
33.
Knight, W. E. and H. W. Benedict. Preliminary report on the
effect of photoperiod and temperature on the flowering and
growth of several southern grasses. Agronomy Journal 45:
268 -269. 1953.
34.
Reproductive activity in Bromus inermis in
relation to phases of tiller development. Botanical Gazette 113:
413 -438. 1952.
35.
Langer, R. H. M. A study of growth in swards of timothy and
meadow fescue. I. Uninterrupted growth. Journal of Agricultural Science 51 :347 -352. 1958.
Lamp, H. F.
134
36.
Control of tillering in grasses by auxin.
American Journal of Botany 36 :437 -439. 1949.
37.
The effect of defoliation, soil fertility,
temperature, and length of day on the growth of some perennial
grasses. Agronomy Journal 37:570 -582. 1945.
38.
Madison, John H.
Leopold, A. C.
Louvorn, R. L.
Turfgrass ecology. Effects of mowing,
irrigation, and nitrogen treatments of Agrostis palustris Huds,
"Seaside" and Agrostis tenuis Sibth, "Highland" on population,
yield, rooting and cover. Agronomy Journal 54(5):407 -412.
1962.
39.
Maeda, Satoshi and Kaoru Ehara. Physiological and ecological
studies on clipping of herbage plants. III. Influence of clipping
on the tiller number and regrowth in Italian ryegrass. Proceedings of the Science Society of Japan 30(4):313 -317. 1962.
40.
Mitchell, K. J. Influence of light and temperature on the growth
of ryegrass (Lolium sp.) I. Pattern of vegetative development.
Physiologia Plantarum 6 :21 -46. .1953.
41.
Mitchell, K. J. Influence of light and temperature on the growth
of ryegrass (Lolium sp. ) II. The control of lateral bud development. Physiologia Plantarum 6:425 -443. 1953.
42.
Mitchell, K. J. Growth of pasture species under controlled
environment. I. Growth at various levels of constant temperature. New Zealand Journal of Science and Technology 38:203216.
1956.
43.
Mitchell, K. J. and R. Lucanus. Growth of pasture species in
controlled environment. II. Growth at low temperature. New
Zealand Journal of Agricultural Research 3:647 -655. 1960.
44.
Mitchell, K. J. and R. Lucanus. Growth of pasture species
under controlled environment. III. Growth at various levels
of constant temperature with 8 and 16 hours of uniform light
per day. New Zealand Journal of Agricultural Research 5:135144.
45.
1962.
Morris, H. S. Physiological effects of boron on wheat. Bulletin of the Torrey Botanical Club 58:1 -30. 1931.
135
46.
Nittler, L. W., T. J. Kenny and Ethel Osborne. Response
of seedlings of varieties of orchardgrass, Dactylis glomerata L.
to photoperiod, light intensity, and temperature. Crop Science
3(2):125 -128.
1963.
47.
Olmsted, Charles E. Growth and development in range grasses.
I. Early development of Bauteloua curtipendula in relation to
water supply. Botanical Gazette 102:499 -519. 1941.
48.
Olmsted, C. E. Growth and development in range grasses. IV.
Photoperiodic responses in twelve geographic strains of side oats grama. Botanical Gazette 106 :46 -74. 1944.
49.
Philipson, W. R. Revision of the British species of Agrostis.
Journal of the Linnean Society of Botany 51:73-151. 1936.
50.
Ryder,
51.
Ryle, G. J. A. and R. H. M. Langer. Studies on the physiology
of flowering of timothy (Phleum pratense L. ). I. Influence of
daylength and temperature on initiation and differentiation of
the inflorescence. Annals of Botany 27(106):213 -231. 1963.
52.
Ryle, G. J. A. A comparison of leaf and tiller growth in seven
perennial grasses as influenced by nitrogen and temperature.
Journal of the British Grassland Society 19(1) :281 -290. 1964.
53.
Shen, T. C. and C. M. Harrison.
On the morphology of
20(4):263 -276. 1954.
V. L.
leaves. Botanical Review
Tillering of Sudangrass
(Sorghum vulgare var. Sudanense Hitsh. ) with special attention
to the effects of ridging. Agronomy Journal 57(5):437 -441.
1965.
54.
Shirley, H. L. The influence of light intensity and light quality
upon the growth of plants. American Journal of Botany 16:354390.
1929.
55.
Silsbury, J. H. Tiller dynamics, growth, and persistency of
Lolium perenne L. and Lolium rigidum Gaud. Australian
Journal of Agricultural Research 15(1):9 -20. 1964.
56.
Smith, R. W. Tillering of grain as related to yield and rainfall.
Journal of the American Society of Agronomy 17:717 -725. 1925.
136
57.
Soper, K. Effects of changes in light and temperature on roots
of perennial ryegrass (Lolium perenne). New Zealand Journal
of Science and Technology 38:783 -792. 1957.
58.
Sprague,
59.
Stapledon, R. G. Cocksfoot grass: Ecotypes in relation to
biotic factor. Journal of Ecology 16:72 -104. 1928.
60.
Stuckey, I. H. Seasonal growth of grass roots.
Journal of Botany 28:486 -491. 1941.
61.
H. B. Root development of perennial grasses and its
relation to soil conditions. Soil Science 36:189 -209. 1933.
Influence of soil temperature on the development
bentgrass. Plant Physiology 17:116 -122. 1942.
Stuckey, I. H.
of colonial
62.
American
Stuckey, I. H. and W. G. Banfield. The morphological variations and the occurrence of aneuploids in some species of
Agrostis in Rhode Island. American Journal of Botany 33:185190.
1946.
63.
Sullivan, J. T. and V. G. Sprague. The effect of temperature
on the growth and composition of the stubble and roots of
perennial rye -grass. Plant Physiology 24:706 -719. 1949.
64.
Warington, Katherine. The influence of length of day on the
response of plants to boron. Annals of Botany 47 :430 -458.
1933.
65.
Wassink, E. C. and J. H. J. Stolwijk. Effects of light quality
on plant growth. Annual Review of Plant Physiology 7:373 -400.
1956.
The growth habits and chemical composition
of bromegrass, Bromus inermis, Leyss, as affected by different environmental conditions. Journal of the American
Society of Agronomy 32:527 -538. 1940.
66.
Watkins, J. M.
67.
Weaver, J. E. J. Kramer and M. Reed. Development of
root and shoot of winter wheat under field environment. Ecology
5:26 -50. 1924.
68.
Weinman, H.
,
grasses.
1948.
Underground development and reserves of
Journal of the British Grassland Society 3:115 -140.
137
69.
Grasses in
agriculture. Rome, Food and Agriculture Organization (of
Whyte, R. O.
,
T. R. G. Moir and J. P. Cooper.
the United Nations),
19 59
.
426 p.
70.
Wood, G. M. and Jane A. Burke. Effect of cutting height
on turf density of Merion, Park, Delta, Newport and common
Kentucky bluegrass. Crop Science 1(5):317 -318. 1961.
71.
Wort, D. J. Soil temperature and growth of Marquis wheat.
Plant Physiology 15:335 -342. 1940.
72.
Youngner, Victor B.
73.
The stooling character of oats.
cultural College Annual Report 38:129. 1912.
Growth and flowering of Zoysia species
in response to temperature, photoperiods, and light intensities.
Crop Science 1(2):91 -93. 1961.
Zavitz,
C. A.
Ontario Agri-
APPENDIX
138
APPENDIX
Appendix Table 1.
Source of
variation
Mean squares and levels of significance for number of tillers, leaf width,
leaf length and diameter of plant. Observation time as a main plot.
Mean squares and significance
Diameter
Leaf
Leaf
Number of
of plant
length
width
tillers
d. f.
BEAVER LODGE
Replic.
3
706.1
.01
16.9
640.5
Obs. time
2
99958.0**
9.02 **
1980.1 **
13596.0**
Error
6
2214.5
.03
63.2
679.5
Clones
4
99208.0**
1.40**
851.4**
2696. 1**
Obs. t. x Cl.
8
21434.0**
32**
395.4 **
1391. 0**
36
1570.9
Error
.
.05
42.2
341.9
SMITHERS
Replic.
3
1901.2
.07
4.2
263.8
Obs. time
2
152400.0**
9.87 **
116.3**
1599.0*
Error
6
3268.9
.08
6.3
152.1
Clones
4
39400. 0**
1. 62**
568. 3**
458.3
Ohs, t. x Cl.
8
5116.9*
.77 **
795.6 **
2083.6
.05
Error
36
1520. 1 **
19.8
275.0
SUMMER LAND
Replic.
3
1751.3
.01
31.3
192.9
Obs, time
2
156020.0**
5.44*
926.5**
27325.0**
Error
6
627.6
.04
86.6
300.8
Clones
4
139360.0*'*
2.39 **
2683.3 **
1050.8*
Obs. t. x Cl.
8
25492. 0**
53 **
599. 4**
826.8*
111.1
360.7
Error
36
2324.7
.
.06
139
Appendix Table 1. (continued)
Mean squares and significance
Number of
Source of
variation
d. f.
tillers
Leaf
Leaf
width
length
Diameter
of plant
244.6
15246.0
256.1
100860.0
254.1
24559.0
VANCOUVER
.1
Replic.
3
3032.5
Obs. time
2
11863.0
Error
6
2536.0
Clones
4
26668. 0**
Obs. t. x Cl.
8
4994. 4*
.6
36
2152.8
.2
Error
5.1 **
.3
1.7 **
*
2947. 1**
186410. 0**
655.2
7403. 3
397.0
8067.5
KLAMATH FALLS
Replic.
3
4250.8
.01
12.9
3668.9
Obs. time
2
154990.0**
15.94**
748.8 **
65540.0 **
Error
6
1945.5
.01
24.2
Clones
4
95378.0**
5.02**
1900.6 **
Obs. t. x Cl.
8
14082.0**
.51 **
134. 1 **
36
1798.7
Error
.04
2335.8
10261.0 **
1109.0
37.7
1199.9
13.5
1976.2
MEDF OR D
Replic.
3
2562.6
.02
Ohs, time
2
29331.0*
19.97 **
Error
6
2904.2
.03
10.7
446.9
Clones
4
90227.0 **
5. 52**
1917. 3 **
25366.0 **
Obs. t. x Cl.
8
14373. 0 **
1. 01**
366. 7**
2865. 3 **
20.8
271.8
Error
36
1052.5
.04
Significant at the five and one percent level respectively.
.8
53892.0 **
140
Appendix Table 2. Mean squares and levels of significance for number of rhizomes, length of
rhizomes and number of rhizome nodes. Observation time as a main plot.
Mean squares and significance
Number of
Source of
Number of
Length of
rhizome nodes
rhizomes
d. f.
hizomes
variation
BEAVER LODGE
Replic.
3
13, 3
588. 7
2, 3
Obs, time
2
1302.1 **
10626.0 **
41.1*
Error
6
27. 6
360, 9
Clones
4
3945. 5**
9069.0 **
Ohs, t. x Cl.
8
1004. 6**
991.
1
9. 1*
19. 6
648.
8
4.0
36
Error
5.
1
44, 3 **
SMITHERS
Replic.
3
52, 2
116.
1
8. 9
2
8656. 2 **
1029, 8
20. 7
Error
6
135. 8
556. 7
4. 5
Clones
4
4075. 4**
3247. 4**
22. 1**
Obs, t, x Cl.
8
3194. 0**
1307. 0*
13.6*
90.6
511.7
4. 9
Ohs.
time
Error
36
SUMMERLAND
36.3
Replic.
3
Ohs, time
2
21615.0 **
8899, 9 **
Error
6
169.9
351.5
3.0
Clones
4
9736. 1 **
2763. 0**
24. 1 **
Ohs, t, x Cl,
8
6018. 2 **
Error
36
123,8
88, 7
,
8
52. 5 **
7
8.6
473, 6
5. 3
736.
141
Appendix Table 2. (continued)
Mean squares and significance
Source of
variation
d. f.
Number of
rhizomes
Length of
rhizomes
Number of
rhizome nodes
VANCOUVER
Replic.
3
569.7
572.6
7.4
Obs. time
2
1221.0
4443.5
16.7*
Error
6
2078.3
1004.9
Clones
4
18346.0**
21036.0 **
65.6**
1
1121.3
7.7
606.5
804.6
4.6
388.
5.2
Obs.
t.
x Cl.
Error
8
36
547.
3.
1
KLAMATH FALLS
Replic.
3
2306.4
Obs. time
2
11758.0*
23525.0 **
138.2 **
Error
6
1804.7
456.6
3.8
Clones
4
35438.0**
Obs. t. x Cl.
8
9257. O**
493.2
6. 5*
2017.0
258.9
2.8
904.2
4.
Error
36
1
3618.5 **
171.7 **
MEDFORD
Replic.
3
440.
Obs. time
2
1270.1
10749.0**
75.8 **
Error
6
368.6
569.2
2.5
Clones
4
14281.0**
10384.0 **
45.2**
Obs. t. x Cl.
8
678, 1**
1965. 7**
11.7 **
Error
36
5
128.1
*, ** Significant at the five and one percent level respectively.
515.1
1
2.8
142
Appendix Table 3. Mean squares and levels of significance for number of stolons, length of
stolons, number of stolon nodes and number of aerial branches.
Observation time as a main plot.
Mean squares and significance
Number of
Length of
Number of
Number of
Source of
aerial branches
stolons
stolon nodes
stolons
d. f.
variation
BEAVER LODGE
R eplic.
3
349.7
2772.4
2.4
.7
Obs. time
2
1771.3
127110.0**
126.1 **
7.9 **
Error
6
349.7
2772.4
2.4
.5
Clones
4
652.7
4148.8
3.8*
8.2 **
Obs. t. x Cl.
8
652.7
4148.8
3. 8 **
7.6 **
36
262.5
2053.7
1.2
.7
.4
2.2
275.6 **
660.0**
Error
SMITH ER
S
65.9
Replic.
3
18.9
Obs. time
2
1215.0**
99349.0**
Error
6
18.9
65.9
.4
2.2
Clones
4
558.4**
16804.0 **
46.4**
69.2 **
8
558.4**
160804.0**
46. 4**
69.2 **
.2
3.5
Obs.
t.
x Cl.
Error
36
11.1
37.8
SUMMER LAND
Replic.
3
668.8
2137.2
1.2
.6
Obs. time
2
58033.0**
344740.0**
344.6**
155.4**
Error
6
666.8
2137.2
1.2
4.0
Clones
4
24572.0**
33567.0 **
36.2**
70.4**
Obs. t. x Cl.
8
24572.0**
33567.0 **
36. 2 **
89.4**
36
409.9
2140.8
1.5
2.5
Error
143
Appendix Table 3. (continued)
Source of
variation
d. f.
Number of
stolons
Mean squares and significance
Length o f
Number o f
stolon nodes
stolons
Number of
aerial branches
VANCOUVER
Replic.
3
349.7
2756.8
2.8
2998.4
Obs. time
2
1771.3
127290.0**
130. 8 **
56747.0 **
Error
6
349.7
2756.8
2. 8
1680.6
Clones
4
652.7
4138.8
3.4*
4800.2*
Obs. t. x Cl.
8
652.7*
4138.8
3.4*
3857.8*
262.5
2058.
3
1.2
1500.6
36
Error
KLAMATH FALLS
Replic.
3
349.7
2653.0
2. 4
640.8
Obs. time
2
1771.3
135850.0**
126. 1 **
151810.0**
Error
6
349.7
2653.0
2.4
623.2
Clones
4
652.7*
3843.9
3. 8*
19068.0 **
8
652.7*
3843.9
3. 8 **
19141.0 **
262.5
2 044. 5
1. 2
153.9
--
965.1
Obs.
t.
x Cl.
36
Error
MEDFORD
Replic.
3
--
Obs. time
2
.-
--
--
428020.0 **
Error
6
--
--
--
1011.6
Clones
4
--
--
--
23989.0 **
Obs. t. x Cl.
8
--
--
--
24569.0 **
36
--
--
434.8
Error
*
**
Significant at the five and one percent level respectively.
144
Appendix Table 4. Mean squares and levels of significance for number of tillers, leaf width,
leaf length and diameter of plant. Location as a main plot.
Mean squares and significance
Leaf
Diameter
Leaf
Number of
Source of
of plant
length
width
tillers
d.
f.
variation
FALL 1964
Replic.
3
2932.9
.04
36.9
4235.9
Location
5
219560.0**
3.62**
1551.0 **
64287.0 **
15
4102.2
.10
85.9
5275.1
4
97640.0 **
7.25 **
3547.2 **
77758.0 **
Loc. x Cl.
20
15341.0**
.52 **
357.0**
12028.0**
Error
72
3123.2
.12
122.0
1064.0
Error
Clones
SPRING 1965
Replic.
3
1522.8
.03
66.4
1380.7
Location
5
16945.0**
18.74**
920.7**
152740. 0**
15
1069.1
.06
50.8
2650.4
4
92282.0**
5.23 **
3142.8 **
39916.0**
Loc. x Cl.
20
1671.1*
.55 **
500.6 **
16551.0 **
Error
72
897,2
Error
Clones
.04
38.0
1134.4
FALL 1965
Replic.
3
2807.6
.04
36.7
7611.2
Location
5
52246.0**
4.47**
1155.9 **
180410.0 **
15
1634,5
.48
29.7
4335.3
4
301030.0**
5.19 **
2109.1 **
44255.0 **
Loc. x Cl.
20
17030.0**
.41 **
770.4 **
10605,0**
Error
72
1472.7
.56
149.5
Error
Clones
*
**
Significant at the five and one percent level respectively.
2558.6
145
Appendix Table 5. Mean squares and levels of significance for number of rhizomes, length of
rhizomes and number of rhizome nodes. Location as a main plot.
Mean squares and significance
Number of
Length of
Number of
Source of
rhizome nodes
rhizomes
rhizomes
d. f.
variation
FALL 1964
Replic.
3
93.7
Location
5
2815. 1**
15
Error
240.9
1849.7
12737.0**
284,9
19.
1
46.3 **
2.
1
4
3423.1**
21402.0*
118.8 **
Loc. x Cl.
20
802. 3 **
1072.6*
6.9
Error
72
57.2
527.0
4.5
81.6
1.3
5667.1 **
17961.0 **
66.9 **
842.8
715.9
5.0
4
32373.0**
15609.0 **
60.3 **
Loc. x Cl.
20
5642.0**
1609.9 **
9.2
Error
72
846.4
734.4
5.8
139.8
.8
Clones
SPRING 1965
Replic.
3
Location
5
15
Error
Clones
946.3
FALL 1965
Replic.
3
1177.3
Location
5
4695. 3 **
1606. 7*
16.6 **
15
593.5
385.6
3.2
4
53686.0**
5087.6 **
27.1 **
Loc. x Cl.
20
2349.5**
1573.0 **
9.2**
Error
72
279.1
360.8
Error
Clones
*
**
Significant at the five and one percent level respectively.
2.1
146
Appendix Table 6. Mean squares and levels of significance for number of stolons, length of
stolon, number of stolon nodes and number of aerial branches. Location
as a main plot.
Mean squares and significance
Length of
Number of
Number of
Number of
Source of
stolons
stolon nodes
aerial branches
d. f.
stolons
variation
FALL 1964
Replic.
3
--
Location
5
--
15
--
113.8
--
--
507. 1 **
--
--
--
99.4
4
--
--
--
260.2 **
Loc. x Cl.
20
--
--
--
158.6 **
Error
72
--
42.6
--
--
Error
Clones
SPRING 1965
Replic.
3
Location
5
--
--
--
--
15
--
--
--
--
4
--
--
--
--
Loc. x Cl.
20
--
--
--
--
Error
72
--
--
-_
Error
Clones
FALL 1965
Replic.
3
469.5
939.4
4.5
Location
5
26146.0**
127870. 0**
129. 9**
208900. 0**
15
528.2
4443.9
9.2
1989.1
4
20740.0**
72843.0**
124.7 **
29929.0**
Loc. x Cl.
20
11326.0**
22363.0**
42. 2 **
22499.0 **
Error
72
342.4
3862.7
5.2
1005.9
Error
Clones
*
**
Significant at the five and one percent level respectively.
698.2
147
Appendix Table 7. Mean squares and levels of significance for number of tillers, amount of
rhizomes, length of rhizomes and number of rhizome nodes. Turf experiment.
Mean squares and significance
Length of
Number of
Amount of
Number of
Source of
rhizome
nodes
rhizomes
rhizomes
tillers
d.
f.
variation
AUGUST 1965
Replic.
7
458.9
7.3
543.4
3.9
Cut, height
3
12702.0**
3.0
1510.4*
9.9
2.8
311.7
5.5
129.2 **
12365.0**
119.7**
5.9 **
961.1
10.9
2.4
680.4
7.7
21
Error
Clones
4
Cut, h. x Cl.
Error
1687.2
19221.0**
12
1124.8
112
667.8
OCTOBER 1965
Replic.
7
564.3
12.6
406.3
1.9
Cut, height
3
4190.2*
112.1
491.3
13.6
1197,9
24.8
475.9
7.0
31038,0**
1071.5 **
11249.0**
115.0**
76.5 **
818.5
10.5
18.3
469.8
6.9
21
Error
4
Clones
Cut. h. x Cl.
Error
12
1244.8
112
781.5
DECEMBER 1965
883.4
12.9
708.8
7.2
8247.4**
31.2*
2149.7
8.1
21
553.6
8.1
800.0
4
54482.0**
438.6 **
12468.0**
12
647.7
12.0
1720.9
20.5
112
582.0
8.3
1143.9
15.5
Replic.
7
Cut, height
3
Error
Clones
Cut. h. x Cl,
Error
* **
Significant at the five and one percent level respectively.
130.3**
148
Appendix Table 8. Mean squares and levels of significance for number of aerial branches,
amount of stolons, length of stolons. and number of stolon nodes. Turf
experiment.
Source of
d. f.
variation
Number of
aerial branches
Mean squares and significance
Length of
Amount of
stolons
stolons
Number of
stolon nodes
AUGUST 1965
R
eplic.
Cut, height
7
449.5
3
50653.0**
21
Error
Clones
Cut, h. x Cl.
1
867.5
2.7
.
09
1575.9
.5
.
11
1455.8
3.8
4
2928.7 **
.12
4384.8
2.7
12
2711.5 **
.14
1648.0
2.4
. 11
1852.8
3.1
7798.9
10.9
112
Error
545.
.12
251.3
OCTOBER 1965
R eplic.
7
12616.0
20.2
Cut, height
3
92071.0**
29.8
6976.7
17.4
21
Error
Clones
Cut, h.
x
Cl.
4718.
1
16649.0**
21.3
7.0
111.5**
4
16800.0**
88.9 **
12
7797. 3 **
18.5*
8549.
8.4
6120.3
9.
112
Error
67912.0**
2114.5
1
6.9
1
DECEMBER 1965
2350.7
9.0
5420.2
14.3
37646.0**
134.9 **
235320.0 **
242.0**
814.2
3.9
6510.7
12.1
4
3357.3 **
235.1 **
5142.6
117.5**
12
589.2
18.6*
7284.6*
7.5
112
437.2
9.1
3258.0
6.5
Replic.
7
Cut. height
3
21
Error
Clones
Cut. h. x Cl.
Error
*
**
Significant at the five and one percent level respectively.
149
Appendix Table 9. Mean squares and levels of significance for number of tillers, leaf width,
leaf length and diameter of plant. Growth chamber experiment.
Mean squares and significance
Diameter
Leaf
Leaf
of
Number
Source of
of plant
length
width
tillers
d. f.
variation
Replic.
7
60.0
Temperature
3
608.0**
Clones
Temp. x Cl.
*
**
2173.5 **
196.1
951.5 **
.06
91.1
170.0
4
4677.3**
4.30**
11303.0**
11276.0**
12
227. 3 **
25 **
205.5 **
345. 8 **
.
.06
53.8
112
Error
.83 **
97.1
91.5
21
Error
.06
75.6
146.2
Significant at the five and one percent level respectively.
Appendix Table 10. Mean squares and level s of significance for number of rhizomes, length
of rhizomes, and number of rhizome nodes. Growth chamber experiment.
Mean squares and significance
Number of
Length of
of
Number
Source of
nodes
rhizome
rhizomes
rhizomes
d. f.
variation
R eplic.
7
.8
598.0
2.8
Temperature
3
47.1**
1376.4
14.9
1.8
1203.5
9.9
4
159.2 **
37897.0 **
205.4**
12
22.0**
1128.2
9.9
2.6
853.1
6.2
21
Error
Clones
Temp. xCl.
Error
*
**
112
Significant at the five and one percent level respectively,