Varieties and Rootstocks – The California Perspective Mary Lu Arpaia and John Menge University of California, Riverside Why Why have have an an Improvement Improvement Program Program Producing the Crop - Enhancing Productivity High production of optimally sized fruit Tolerance to disease, pests Tolerance to environmental stress Tree "manageability" On-tree storage and minimize alternate bearing Marketing the Crop - Maximizing Fruit Quality Minimizing physiological disorders Uniformity of ripening Tolerance to low storage temperatures Tolerance to handling "mis-management" Decreased postharvest fruit decay Mary Lu Arpaia (September 2004) 1 Requirements/Risks of a plant improvement program Long-term venture (10 – 20 years) 9 Requires coordinated effort: academia, growers, packers, consumers 9 Wide-scale adoption unknown: will there be a return on the investment? 9 Most current cultivars and rootstocks are local selections – can improvements be found? 9 Challenges Challenges Specific Specific to to Avocado Avocado Plant Plant Improvement Improvement Avocado is “relatively primitive” – commercial production <100 years 9 Understanding limited on avocado genetics 9 May be many years to come into production 9 Highly heterozygous – seedling populations extremely varied 9 Mary Lu Arpaia (September 2004) 2 The goals of an avocado improvement program can be achieved through varietal and rootstock manipulation using either traditional breeding methods or as technology improves, molecular techniques. Additionally, an important component for the future is germplasm conservation. Characterization and preservation of wild Persea and related genera is essential in order to preserve desirable traits useful for future breeding efforts. University University of of California California Avocado Avocado Improvement Improvement Program Program Rootstock Selection Program Disease tolerance - J. Menge, G. Zentmyer Salinity tolerance - D. Crowley, M. L. Arpaia Field Productivity - M. L. Arpaia, G. Bender, B. Faber Varietal Improvement - M. L. Arpaia Genetic Characterization - M. Clegg, T. Chao Germplasm Conservation - R. Scora, J. Menge, M. L. Arpaia Mary Lu Arpaia (September 2004) 3 Contributions Contributions of of UC UC Avocado Avocado Program Program Rootstocks Identification of PRR tolerant material Rootstock productivity and salinity tolerance Dwarfing Cultivars New varieties for CA growers Breeding stock shared with international community and has been the foundation of other intl. breeding programs Components • Testing current selections • Developing new selections Mary Lu Arpaia (September 2004) 4 Varietal Selection UC, Riverside B. O. Bergh: 1964 - 1994 G. W. Witney: 1994 - 1996 M. L. Arpaia: 1996 - present Technical Support: Bob Whitsell 1964 - 1989 Gray Martin 1984 – 1997 David Stottlemyer 1997 - present Varietal Varietal breeding breeding in in California California Current major cultivars are "local" or introduced selections Released Cultivars to CA industry Dr. B. O. Bergh and Mr. R. Whitsell Gwen, Whitsell and Esther - released in 1984 Dr. B. O. Bergh and Mr. G. E. Martin Lamb Hass and Sir Prize - released in 1996 Released 2003 Dr. Mary Lu Arpaia and Mr. David Stottlemyer 3-29-5 (GEM) and N4 (-) 5 (Harvest) Mary Lu Arpaia (September 2004) 5 Components Components of of evaluation evaluation • • • • • Yield Fruit characteristics – size, seed size Maturity and postharvest quality Tree vigor – growth habit Flowering, stress tolerance Mary Lu Arpaia (September 2004) 6 Growth Habit Varies Hass 3=29-5 (GEM) Gwen Lamb Hass N4 (-) 5 (Harvest) Marvel (BL516) Leaf Shape - The unreleased varieties are all ‘Gwen’ offspring. Although the leaves tend to be similar, there are subtle differences between the varieties. Mary Lu Arpaia (September 2004) 7 Yield UC South Coast REC, Topworked Trees 1600 1600 1400 1400 Fruit Count Fruit Count 1200 1200 1000 1000 2004 2004 2003 2003 2002 2002 2001 2001 2000 2000 1999 1999 800 800 600 600 400 400 200 200 0 0 OA184 OA184 BL667 BL667 (Nobel) (Nobel) BL516 BL516 (Marvel) (Marvel) Hass Hass Harvest GEM Harvest GEM (N4 (-) 5) (3-29-5) (N4 (-) 5) (3-29-5) Sir Prize Lamb Hass Sir Prize Lamb Hass Cultivar Cultivar Alternate Bearing Tendency UC South Coast REC, Topworked Trees Less More Lamb Hass Sir Prize GEM (3-29-5) Harvest (N4 (-) 5) Hass BL516 (Marvel) BL667 (Nobel) OA184 0 25 50 75 100 125 150 175 200 Coefficient of Variation (%) Mary Lu Arpaia (September 2004) 8 Yield DeBusschere Variety, Clonal Duke 7 RS Fruit Fruitcount Coun 900 800 2004 700 2003 600 2001 2002 500 400 300 200 100 0 BL667 BL516 Lamb Hass Harvest Hass GEM Cultivar Flowering at UC South Coast REC in Irvine, CA. for 2002 - 2004 Hass Lamb GEM Harvest Sir Prize BL667 BL516 0 20 40 60 80 100 120 140 160 180 Julian Date 2004 2003 2002 Flowering varies from year to year Mary Lu Arpaia (September 2004) 9 Fruit Maturity 3-29-5 (GEM) 40 40 GEM Hass 2000 2002 2003 35 Dry Matter Content (%) Dry Matter Content (%) 35 2001 30 25 20 30 25 20 15 15 1/1 1/15 1/29 2/12 2/26 3/11 3/25 4/8 4/22 5/6 5/20 6/3 6/17 7/1 7/15 7/29 8/12 8/26 9/9 9/23 1/1 1/15 1/29 2/12 2/26 3/11 3/25 4/8 4/22 5/6 5/20 6/3 6/17 7/1 7/15 7/29 8/12 8/26 9/9 9/23 10/710/21 2003 Calendar Date Similar pattern of dry matter accumulation to Hass Comparison of dry matter changes over season Fruit Maturity N4 (-) 5 (Harvest) 35 40 Harvest Hass 2000 2001 2002 2003 30 Dry Matter Content (%) Dry Matter Content (%) 35 25 30 25 20 20 15 15 1/1 1/15 1/29 2/12 2/26 3/11 3/25 4/8 4/22 5/6 5/20 6/3 6/17 7/1 7/15 7/29 8/12 8/26 9/9 9/23 2003 Slower pattern of dry matter accumulation to Hass Mary Lu Arpaia (September 2004) 1/1 1/15 1/29 2/12 2/26 3/11 3/25 4/8 4/22 5/6 5/20 6/3 6/17 7/1 7/15 7/29 8/12 8/26 9/9 9/23 10/7 10/21 Calendar Date Comparison of dry matter changes over season 10 Differences between Hass and Lamb Hass Lamb Hass maturity season – mid to late summer Fruit shape – more “square” Lamb Hass has more upright growth habit Flexible wood – fruit borne interior of tree; tends to set fruit in clusters Lamb Hass is more “tolerant” to Persea mite and other pests (?) Photosynthetic rate approximately 30% higher than Hass and higher chlorophyll content Fruit Maturity – Lamb Hass 40 % Dry Weight 36 Hass '99 Lamb '99 Hass '00 Lamb '00 Hass '01 Lamb '01 32 28 24 20 16 Jan 1 Jan 29 Feb 26 Mar 26 Apr 23 May 21 Jun 18 Jul 16 Aug 13 Sep 10 Delayed dry matter accumulation compared to Hass Pattern of accumulation similar over seasons Mary Lu Arpaia (September 2004) 11 Growth habit differences between Hass and Lamb Hass Hass Lamb Hass Developing new selections 1. In consultation with U. Lavi (Israel) Seed collection: BL667, BL516, GEM, Gwen, Lamb Hass, 5-552 2. In consultation with B. Bergh Isolation Blocks: BL516 x GEM Gwen self Thille x GEM Lamb x GEM Sir Prize x Gwen BL667 x Lamb Mary Lu Arpaia (September 2004) 12 Indexing for disease is a critical component Critical to know: •Relative disease tolerance •All introduced material should be tested Clonal Rootstocks The California Experience Mary Lu Arpaia (September 2004) 13 Rootstocks can influence many scion characteristics Yield Tree size/vigor Yield efficiency Leaf nutrient status Tolerance to environmental stresses • • • • • Use of clonal rootstocks relatively new • • • 500 Potential for future improvements high Significant differences due exist More uniform tree performance possible Cumulative yield per tree (kg) 2 Years after Planting 4 5 6 7 8 3 9 10 400 300 200 100 0 Borchard 100 Duke 7 Toro Canyon Topa Topa D9 G755A G755B G755C Canopy volume (cu. meters) Borchard Duke 7 Toro Canyon Topa Topa D9 G755A G755B G755C 80 60 40 Rootstock influences yield and tree size 20 4.5 6.5 7.5 8.5 Years after planting Mary Lu Arpaia (September 2004) 14 Tree Size – South Africa Duke 7 2 G755C Yield efficiency per cycle (kg/cu. m canopy) 1.5 Duke 7 Toro Canyon 1 D9 G755C 0.5 0 Year 4-5 Yield efficiency declines due to increases in tree size and loss of productive canopy surface Year 6-7 Year 8-9 Table 6 Yield efficiency (kg fruit·m-3) of ‘Hass’ avocado trees growing on ten clonal rootstocks at the University of California South Coast Research and Education Center (latitude, 33°44’N; longitude, 117°49’W) (n=20). Yield efficiency (kg fruit·m-3) Rootstock Significant differences in yield efficiency Not necessarily associated with vigorous trees Mary Lu Arpaia (September 2004) 1991/92 1993/94 1995/96 mean G755A G755B G755C Topa Topa Duke 7 Borchard D9 Toro Canyon 0.60c 0.48cd 0.42d 0.81b 0.97a 0.96ab 0.95ab 0.90ab 0.87c 0.83cd 0.69d 1.06b 1.23a 1.17ab 1.15ab 1.13ab 0.238c 0.182c 0.236c 0.935a 0.958a 0.935a 0.696b 0.948a 0.57c 0.50cd 0.45d 0.93b 1.05a 1.02ab 0.93b 0.99ab Thomas G1033 ∗∗∗ 0.75 0.58 ∗∗∗ 1.00 0.94 n.s. ∗∗∗ 0.50 0.69 ∗∗∗ 0.75 0.74 n.s. ∗ ∗ Mean separation tests within columns non-significant (n.s.), or significant at the P=0.05 (∗) or 0.001 (∗∗∗) level based on Fisher’s Protected LSD test. 15 Alternate Bearing can be influenced by clonal rootstock 140 Weight per tree (kg) Duke 7 120 Toro Canyon 100 D9 80 G755C 60 40 20 0 2 ** 3 ** 4 ** 5 ** 6 ** 7 ** 8 * 9 ** 10 ** Years after planting Table 2. Alternate bearing index of ‘Hass’ avocado trees growing on ten clonal rootstocks at the University of California South Coast Research and Education Center (latitude, 33°44’N; longitude, 117°49’W). See Materials & Methods section for calculation. Significant differences detected Rootstock Alternate bearing index G755A G755B G755C Topa Topa Duke 7 Borchard D9 Toro Canyon 1.49c 1.25d 1.21d 1.91a 1.61bc 1.70b 1.56bc 1.92a Thomas G1033 ∗∗∗ 1.83 1.63 n.s. Mean separation tests within columns non-significant (n.s.) or significant at the P=0.001 (∗∗∗) level based on Fisher’s Protected LSD test. Mary Lu Arpaia (September 2004) 16 Rootstock can influence leaf nutrient status Nitrogen Phosphorus 2.7 0.255 0.235 2.5 0.215 0.195 % dry weight % dry weight 2.3 G755C Duke 7 2.1 D9 Toro Canyon 1.9 G755C 0.175 Duke 7 0.155 D9 Toro Canyon 0.135 0.115 1.7 0.095 0.075 1.5 1988 1989 1990 1991 1992 1993 1994 1988 1995 1989 1990 1991 1992 1993 1994 1995 Year Year Potassium 1.7 1.6 % dry weight 1.5 G755C 1.4 Duke 7 1.3 D9 Toro Canyon 1.2 1.1 1 1988 1989 1990 1991 1992 1993 1994 1995 Rootstocks affect ‘Hass’ avocado fruit rots – believed related to calcium uptake 4 wks @ 5C Flesh volume (%) Marques, Hofman et al 2001 40 SV Seedling Velvick CV Clonal Velvick D7 Duke 7 30 b a c 20 a b 10 a Non-stored Mary Lu Arpaia (September 2004) Stored 17 ‘Hass’ Avocado Phenology Model Relative growth rate “On” year = heavy crop load “Off” year = light crop load 100 80 shoot 60 root 40 bloom 20 0 Julian date Are there rootstocks affects? Root growth rate – No consistent differences Annual root growth rate of ‘Hass’ avocado on four clonal rootstocks. Year Rootstock 1992 1993 1994 -1 mm·day ·root ‘Thomas’ ‘Topa Topa’ ‘Duke 7’ ‘D9’ Sig. of Fx 0.75ab 0.84a 0.60c 0.67bc * 0.72 0.97 0.69 0.72 0.67a 0.75a 0.47b 0.75a n.s. * 1995 1996 0.52a 0.32bc 0.18c 0.38ab 0.38a 0.25b 0.23b 0.24b -1 ** * z Means within a column with no letter(s) in common are significantly different (Fisher’s Protected Least Significant Difference test at P=0.05). y ns, *, **, *** are non-significant, or significant at P≤0.05, P≤0.01, or P≤0.001, respectively. Mary Lu Arpaia (September 2004) 18 Rootstock had no effect of shoot growth rate Shoot growth ratez (mm·day-1·shoot-1) of ‘Hass’ avocado on one of four rootstocks. ‘Thomas’ ‘Topa Topa’ ‘Duke 7’ ‘D9’ Sig. of Fy Avg. 1992 1993 1994 1995 1996 springz summer spring summer spring summer spring summer spring summer 0.68 0.65 0.61 0.63 0.37 0.26 0.27 0.16 3.57 5.71 5.34 5.05 4.57 5.81 6.49 6.59 1.60 1.80 2.00 1.82 0.46 0.36 0.46 0.72 0.83 0.51 0.70 0.86 0.78 0.41 0.92 0.46 1.40 0.97 1.17 1.34 0.37 0.20 0.32 0.26 n.s. n.s. n.s. n.s. n.s. n.s. n.s. n.s. n.s. n.s. 0.64 0.27 4.92 5.86 1.80 0.50 0.73 0.64 1.22 0.29 z Spring = average of spring (first) flush; Summer = average of summer (second) flush. y ns, *, **, *** are non-significant, or significantly different at P≤0.05, P≤0.01, or P≤0.001, respectively. Statistical analysis performed using log transformed growth rates (log10 of rate + 1). Environmental Stress – Phytophthora root rot Disease resistance/tolerance possible New selections Young avocado with root rot resistant rootstock growing among older avocados dying from avocado root rot Mary Lu Arpaia (September 2004) 19 Breeding program Breeding blocks of resistant varieties planted together to enhance natural crossing. Screening and greenhouse evaluation of rootstocks Fruit from breeding blocks is germinated in the greenhouse and inoculated with Phytophthora cinnamomi Mary Lu Arpaia (September 2004) Selected resistant seedlings are reinoculated each time they are repotted 20 Resistant varieties are grafted to stumps in the field to get abundant budwood for experiments. Production of clonal rootstocks for experiments Field Evaluation Thomas Steddom Resistant rootstocks grown in Phytophthora-infested soil Uzi Mary Lu Arpaia (September 2004) Thomas 21 Environmental Stress – Salinity San Diego County Stehly Trial Duke 7 Latas Toro Canyon VC801 VC218 Thomas PP4 VC256 Zutano December 2001 Mary Lu Arpaia (September 2004) 22 Condition of surviving trees - July 2002 5.0 4.5 4.0 3.5 3.0 2.5 2.0 1.5 25 VC 2 4 yo n PP To ro Ca n 5 PP Th om as * 18 VC 2 7 41 Du ke VC 2 01 56 VC 2 Ev st ro VC 8 ta no uk e* D Ba rr Zu La ta s 1.0 Chloride concentration - September 2001 Sodium concentration - September 2001 350.0 0.14 300.0 0.12 250.0 0.10 200.0 0.08 150.0 0.06 100.0 0.04 50.0 0.02 0.00 0.0 Latas Zutano Barr Evstro VC801 VC256 Duke 7 VC241* VC218 Duke* PP5 Thomas PP4 Toro Canyon VC225 Latas Zutano Barr Evstro VC801 VC256 Duke 7 VC241* VC218 PP5 Thomas PP4 Duke* Toro VC225 Canyon What we know - Much yet to be learned Differences between avocado races needs to be better understood Clonals can improve overall tree performance and potentially postharvest quality As with other tree crops; with time we will find that varying environmental conditions will require different rootstocks • Salinity, Disease, Cold, Soil Mary Lu Arpaia (September 2004) 23 Greater cultural, harvesting and water costs coupled with increasing market competition Approaches for the future Enhancing Productivity Understanding avocado tree physiology and stress responses • Light manipulation • High density plantings • Rootstocks for disease, salinity tolerance, dwarfing • Varieties w/ > productivity, pest tolerance, suitable for close spacings • Pollinizers and Pollinators • NEED INTEGRATED PROGRAM For For more more information information www.avocadosource.com www.avocadosource.com Variety Variety information information •• Database Database of of avocado avocado varieties varieties •• Information Information on on rootstocks rootstocks (ongoing) (ongoing) •• Mary Lu Arpaia (September 2004) 24