Fruit Flies Play Role in Sour Rot Complex: Training system can impede development of disease.
In many other parts of the world, and in most of the published literature, sour rot is a syndrome characterized by decay, oxidation (browning) of the berry skin and the smell of acetic acid (vinegar) emanating from diseased grapes that break down and begin to leak their contents, hence the "sour" designation.
Although mold fungi such as those in the California complex are sometimes present as secondary decay organisms, they are not necessary components of this form of sour rot elsewhere (see photo above). Studies conducted on Riesling, Pinot Noir and other V. vitiifera cultivars in Ontario, Canada, have shown that they are not susceptible to such disease development until they reach a maturity stage of 15[degrees] Brix.
To successfully manage a disease, it is important to identify the specific organisms involved and how they interact with the plant host and environment to produce the condition. In the past four years, we have studied sour rot in the laboratory and a series of vineyard trials in New York to identify specific organisms and conditions that are needed for sour rot to develop there and, consequently, how to manage it. This has allowed us to develop a definition of the disease as seen in eastern North America, Europe, Oregon and elsewhere--as well as a pesticide program to manage it--and to identify significant differences in disease severity related to trellising systems.
Wounds are important
Although sour rot occurs sporadically, it has been reported worldwide where and when conditions for the disease are favorable. Such conditions are poorly defined but often include pre-harvest rains and temperatures regularly exceeding 65[degrees]F during the final stage of ripening. Once sour rot begins, typically following rain that occurs after berries reach 15[degrees] Brix, it often spreads rapidly.
Wounds are necessary for entry of the yeast, bacteria and fruit flies that are causal components of the sour rot complex, and these microorganisms are distributed to them by rain or heavy dew. Although such wounds are sometimes due to birds and insects, sour rot more commonly begins through cracks in the grape skin caused by rain or by swelling of the berries as they take up water and pull away from pedicels (berry stems) in tightly compressed clusters shortly before harvest.
In the summer bunch rot complex in California, rain is not a requirement. Skin-splitting from irrigation, temperature flux and/ or rapid growth appear to provide wounds for entry of the causal organisms, then wind, insects and perhaps birds seem to facilitate disease spread. It also seems likely that the various mold fungi that enter these wounds as early as 8[degrees] Brix cause decay and further breakdown of the berries, allowing yeast and bacteria to enter later and produce the acetic acid responsible for the smell of vinegar, although this has not been proven.
Wounds also let oxygen into the berry. This is necessary for the conversion of ethanol, first produced by yeasts, to acetic acid by specific bacteria within the berry. In turn, volatilization of ethanol and acetic acid attract Drosophila fruit flies that feed on the yeast and bacteria producing these compounds, and the insects then spread the microorganisms to new berries.
Determining the cause of sour rot
Our study began with the collection of sour-rotted clusters from 16 vineyards in the Finger Lakes region of New York. High levels of both ethanol and acetic acid were found (averaging 1.14 g/L and 1.58 g/L, respectively, in two years of sampling). More than 1,300 individual microbial isolates were recovered, more than 90% of which fell into four categories: two Metschnikowia spp. yeast strains, Gluconobacter cerinus (an acetic acid bacterium, or AAB), and a ubiquitous environmental bacterium (Rahnella sp.) neither associated with acetic acid production nor considered a plant pathogen.
A series of experiments was begun in the lab to determine whether the recovered microorganisms were, in fact, causing the disease. Berries of V. vinifera cultivars Cabernet Franc, Chardonnay and Red Globe were wounded and inoculated with 21 different individual yeasts, mold fungi, bacteria or combinations thereof. One set of inoculated berries was exposed to fruit flies (Drosophila melanogaster), while a companion set was not. After eight days of incubation, berries were rated for rot and browning on a scale of 0-4, then macerated and the juice measured for acetic acid content. Berries were determined to have sour rot if they had a visual disease rating of 3 or 4 and an acetic acid content of at least 0.83 g/L (based on the ranges detected in field samples).
Across all three cultivars, the only combinations of organisms that caused sour rot per our criteria were:
* Aspergillus niger (filamentous fungus) x Gluconobacter oxydans (AAB)
* Hanseniaspora uvarum (yeast) x Acetobacter aceti (AAB)
* Hanseniaspora uvarum x Acetobacter aceti x Gluconobacter oxydans
* Pichia kluyveri (yeast) x Acetobacter aceti
* Pichia kluyveri x Gluconobacter oxydans
* Saccharomyces cerevisiae (yeast) x Acetobacter aceti
* Saccharomyces cerevisiae x Gluconobacter oxydans However, these combinations produced sour rot symptoms only on berries also exposed to wild type Drosophila fruit flies, which carry their own microbiota. When inoculated berries were not exposed to the flies, typical sour rot symptoms did not develop.
To determine whether the effect of the flies was due to microorganisms that they contributed or to some non-microbiological factor, colonies of axenic fruit flies were developed. These were reared in sterile media, rendering them devoid of gut and surface microbiota. In a series of similar inoculation experiments utilizing Red Globe grapes and multiple combinations of microbes plus axenic fruit flies (or not), only the following microbial combinations (each consisting of a yeast and AAB species) consistently caused sour rot symptoms, and only in the presence of axenic flies:
* Saccharomyces cerevisiae x Acetobacter aceti
* Saccharomyces cerevisiae x Gluconobacter oxydans
* Pichia kluyveri x Acetobacter aceti
* Pichia kluyveri x Gluconobacter oxydans
Three components essential for sour rot disease development
The results show that three components are essential for sour rot disease development:
Yeast, which first produce ethanol from the juice of affected grapes,
Acetic acid bacteria, which convert this ethanol to acetic acid,
Drosophila fruit flies, which contribute to the breakdown of infected berries and the liberation of acetic acid volatiles responsible for the characteristic vinegar smell. (Similar results were obtained in some parallel studies comparing the effects of D. melanogaster, the common fruit fly, and D. suzukii, the spotted wing Drosophila, which is not a common pest on grapes in North America). None of these elements cause disease symptoms on their own.
In nature, yeast and acetic acid bacteria may originate from several sources: the outer surface of healthy berries (albeit in low numbers), inside healthy berries (various non-Saccharomyces yeasts such as Pichia spp. were repeatedly isolated from inside healthy berries obtained from vineyards in New York, California and Washington state, in addition to AAB on occasion) and Drosophila-fruit flies.
In addition to making a vital non-microbial contribution to sour rot development by facilitating their breakdown, fruit flies also vector the causal yeast and bacteria both on the outside of their bodies and by transferring gut microbes during feeding, thereby spreading the disease rapidly once it has begun. Thus, sour rot control programs in the vineyard ideally should include measures targeting both the abovementioned microbes and fruit flies.
Chemical control trials
In three spray trials conducted in 2013, 2015 and 2016 in Geneva, N.Y., various antimicrobial and insecticide treatments-both alone and in combination-were applied to a vineyard planted to Vignoles (a tight-cluster interspecific hybrid variety) to test their effect on sour rot development.
In each trial, alternate vine rows were sprayed with an insecticide (Delegate in 2013 or Mustang Maxx in 2015 and 2016) or left untreated. One- or two-panel plots were treated with antimicrobial materials (potassium metabisulfite, Kocide 3000, OxiDate 2.0 or Fracture) at various timings and rates (see treatment list and results in the table "Control of Sour Rot with Anti-Microbial Sprays +/- Insecticide" on page 124). Note that potassium metabisulfite (KMS), a common disinfectant in the winery, is not registered for use in the field but was included in these tests as a "proof of concept."
Insecticides were applied weekly when fruit reached 15.0[degrees] Brix, the stage where berries start to become susceptible to sour rot. The start of weekly antimicrobial treatments varied; their timing is listed in the table as:
* Pre-symptoms starting at 15[degrees] Brix,
* Starting when sour rot symptoms were first visible,
* After the first rain following 15.0[degrees] Brix,
* Following an increase in maximum daily dew point over three days, and
* No treatment.
Insecticide (plus antimicrobials) provided control
In 2013, antimicrobial and insecticide treatments applied in combination provided significant control of sour rot with a reduction in disease severity (percent of cluster area diseased) of 30% to 54% compared to the unsprayed treatment. The vines treated solely with antimicrobial sprays did not experience a significant reduction in sour rot, nor did the treatment in which only insecticide was applied.
Significantly greater sour rot pressure was seen in 2015, with nearly 30% of berries diseased on vines receiving no insecticide or antimicrobial treatment, compared to 16% in 2013. In contrast to 2013, the insecticide alone provided significant control (57% reduction relative to untreated vines) without addition of an antimicrobial treatment. Disease control increased significantly (73% to 84% reduction) in those panels where both antimicrobial and insecticide treatments were applied weekly starting before the onset of symptoms.
However, when the addition of antimicrobial treatments was delayed until symptoms appeared, the combination treatments did not provide significantly more control than the insecticide alone. Furthermore, even when applied on a preventive basis before symptoms appeared, antimicrobial treatments applied alone were relatively ineffective and never provided the same level of control as insecticide applied alone.
Sour rot severity on untreated vines in 2016 was comparable to 2015. Significant control (40%) was provided when insecticide was applied without the addition of an antimicrobial treatment. Further control was achieved when any of three antimicrobial treatments was combined with insecticide starting before the onset of symptoms, whereas delaying addition of an antimicrobial treatment until symptom onset increased control for only one of the two materials tested (see "Sour Rot Control, Field Trial," above).
Three years of chemical control trials demonstrated the importance of insecticide sprays in controlling sour rot, and the additional control provided when antimicrobial sprays were combined with insecticide before the onset of symptoms. In all three years, applying KMS weekly beginning at 15[degrees] Brix (pre-symptoms) in conjunction with an insecticide achieved an average 65% control compared to the unsprayed treatment, and in the two years when OxiDate 2.0 was included in conjunction with Mustang Maxx on this schedule, an average 69% control was achieved. In the two years when Mustang Maxx was the insecticide used, it provided significant control (48% average) when applied alone, whereas antimicrobial treatments applied alone on this same pre-symptom schedule provided only 28% control when averaged for all materials across the three trial years. For growers deciding whether to apply only an insecticide or antimicrobial product, the insecticide appears to be the more important component of the mix.
Also noteworthy is that initiating antimicrobial sprays before the onset of symptoms was more effective than using a limited number of applications after symptoms appeared. The experimental design did not allow examination of the effect of insecticide applications alone if initiated after development of disease symptoms.
Training system effects
In a commercial vineyard of Vitis interspecific hybrid Vignoles in Branchport, N.Y., one block was divided into 14 vine rows trained in a vertical shoot position (VSP) system and 14 vine rows trained to a high-wire cordon (HW) system. Significant sour rot severity was observed in the vineyard block in previous years. The goal was to study whether or not the training system had a significant effect on disease development.
In 2014-16, one vine was selected in each of 20 rows (10 in the VSP section and 10 in the HW section of the block). Following the first sighting of sour rot symptoms in the vineyard, disease ratings were made every three to four days until harvest.
In all three years of the study, there was significantly more sour rot at harvest in vines trained to the HW trellis system than the VSP system. In 2014, for example, sour rot severity increased from 21% to 35% in the HW system and from 13% to 18% in the VSP system in the final seven days before harvest (see "Effect of Training System on Sour Rot Development," above).
This study served not only to examine the differences between training systems but also to document the rapid progression of disease severity as it increased steadily in both training systems over the final one to two weeks before harvest. Measurements of canopy density showed HW vines to have significantly greater density than VSP vines between the vineyard floor and fruiting zone, due to the umbrella-like structure provided by their drooping shoots. This likely provided less ventilation and a more favorable microclimate for disease than the VSP system, where vines were denser above the trellis wires due to upright growth of the shoots and their retention by catch wires.
Integrated control is necessary
The sour rot complex is a dynamic system involving yeast, AAB and Drosophila fruit flies. The species of yeast can vary, as it can for the AAB and fruit flies, yet all three components must be present in order for symptoms to develop. An integrated control program for sour rot should utilize both canopy management and spray applications that target yeasts, AAB and Drosophila spp., although the best protocol for timing of such spray applications has yet to be determined.
Additional trials under commercial or large-plot conditions should help identify the most efficient timings to balance the competing desires to minimize spray applications while maximizing control, although existing data suggest that control is likely to be maximized by initiating sprays before an epidemic is in progress.
It is important to note that these studies have been conducted with a susceptible cultivar in a climate favorable to development of severe disease symptoms. While we did not specifically study cultivars and climatic variables, management recommendations may vary for vineyards with cultivars that have thicker skins and looser clusters and/or in a less conducive environment for the disease.
Nevertheless, this perspective on understanding the sour rot complex is not only applicable to New York grapegrowers but may be pertinent to growers worldwide who experience sour rot as a challenge in their vineyards. The sour rot complex is somewhat unique because we are targeting multiple unrelated causal organisms in control programs, but now there is a more comprehensive understanding of how the disease develops and, in turn, how to manage it.
Although sour rot is sometimes used as a catch-all term to refer to a complex of late-season bunch rots that develop in tight-cluster or thin-skin varieties caused by numerous fungi, the authors have defined it more narrowly as a condition involving decayed berries with brown (oxidized) skins often free of fungal contaminants and with pulp that smells of vinegar (acetic acid).
Sour rot-affected berries collected from numerous vineyards contained significant amounts of ethanol (produced by various yeast species) and acetic acid (produced by specific bacteria) and were always associated with large populations of Drosophila fruit flies.
Experimental inoculation in the laboratory with microbes recovered from diseased clusters was not sufficient to reproduce sour rot symptoms unless inoculated berries also were exposed to Drosophila fruit flies.
Sour rot results from the interaction of yeasts that ferment grape pulp to ethanol, specific bacteria that oxidize ethanol to acetic acid and Drosophila fruit flies, which appear to be responsible for the breakdown of infected berries and subsequent spread of the disease.
Three years of vineyard spray trials with antimicrobial treatments targeting the causal yeast and bacteria alone achieved modest reductions in sour rot severity.
Insecticides targeting Drosophila fruit flies significantly reduced sour rot severity, and combining antimicrobial sprays with the insecticide improved it even further.
Mid-wire cordon-trained Vignoles grapevines had less sour rot when trained with vertical shoot positioning than when trained with a high-wire cordon system where drooping shoots enclosed the fruit within an umbrella-like canopy.
Management of sour rot involves controlling both the microbes and the Drosophila fruit flies that cause the disease and is aided by viticultural practices that enhance ventilation and exposure of the fruit.
This text was adapted from the original version, which appeared in Appellation Cornell, a quarterly newsletter published by Cornell University.
HIGH-WIRE CORDON TRAINING SYSTEM
Current-year shoots droop downward from their origin on the permanent cordon, 5.5 feet above the ground.
VSP TRAINING SYSTEM
Current-year shoots rise from the fruiting-wire cane 3.5 feet above the ground and are maintained in a vertical position by the catch wires above.
Source: Horticulture Section, School of Integrative Plant Science, Cornell University.
Related Article: SUMMER BUNCH ROT (SOUR ROT) IN CALIFORNIA.
By Paul Verdegaal
Zinfandel growers in California are accustomed to being on the lookout for sour rot (summer bunch rot) in any given year, and 2017 was such a growing season. Because of the many factors and pests involved in sour rot, the key to controlling it is still not well understood. Summer bunch rot develops after bunch closure, as berries begin veraison and sugar content exceeds 8[degrees] Brix (from personal observation, it seems to be about 75% of clusters with any color or softening, even one berry). It is hard to predict when, where and how much of a problem sour rot can be.
Any injury or break in berry skins make clusters susceptible to invasion by a wide variety of fungi. Depending on region, variety and site, the range of fungi and bacteria can be extensive but usually includes yeast species present, Acetobacter spp. and opportunistic molds such as Aspergillus niger, Alternaria carbonarius, Alternaria tenuis, Cladosporium herbarum, Rhizopus arrhizus Penicillium sp., Botrytis cinerea at times and others.
With veraison, summer bunch rot becomes apparent with the odor of vinegar and the signs of "leakers" (clusters leaking juice on the vineyard floor). Ultimately, berries and clusters can dry and shrivel to a mass of rot infested with Drosophila fruit flies. As in nearly all other parts of North America where studies have been conducted, these are the common fruit fly, D. melanogaster, not the spotted wing drosophila (D. sizukii), which seems not to be a wine grape pest in California. Growing conditions are one factor, with actual vine growth and fruit development resulting in thin skins or berry cracking, management of water and nutrients, or damaging agents such as powdery mildew, Botrytis, insects, birds, etc.
Tight clusters and varieties with thin skin are most susceptible, with Zinfandel most often affected in Lodi, Calif. Other varieties such as White Riesling and Petite Sirah can have the problem, along with Sauvignon Blanc and Pinot Grigio or old stalwarts like Carignan.
In 2017, high rainfall in Lodi led to vigorous and rapid growth of shoots and fruit. Powdery mildew pressure, alternate periods of hot and cool weather and windy days, along with variable soil moisture as soils dried and required irrigation during heat waves, may have all combined to weaken skins. Sour rot followed in late July, later than normal.
Research indicates some mitigation with preventive sprays including Switch, Pristine and maybe copper (see University of California IPM Guidelines). But these are only slightly helpful and, for the cost, most often not a satisfactory strategy. Careful management of irrigation and nutrients, crop load and shoot growth of the canopy are the best that can be done.
The 2017 season was difficult in Lodi in many aspects, and sour rot (especially in Zinfandel) was fairly severe.
Paul Verdegaal is a University of California farm advisor for San Joaquin County.
OBSERVATIONS FROM THE PAST 20 YEARS Year Sour Rot Level 2016 Very little 2015 Scattered light 2014 Moderate 2013 None 2012 Light 2011 Severe (and Botrytis) 2010 Light (with berry shrivel) 2009 Light (mostly Botrytis) 2008 Slight 2007 None 2006 Severe 2005 None 2004 None 2003 Light (Botrytis bad) 2002 Moderate 2001 None 2000 None 1999 None 1998 Light (Botrytis) 1997 None Periods of alternating hot and cool weather can weaken the skins of wine grapes, promoting sour rot.
Megan Hall was a graduate research assistant at Cornell University's New York State Agricultural Experiment Station and is now an assistant professor of viticulture at the University of Missouri. Gregory Loeb is a professor in the Department of Entomology at Cornell University. Wayne Wilcox is a plant pathology and plant-microbe biology professor at Cornell University.
The authors extend special thanks to Finger Lakes grapegrowers for allowing the use of their vineyards for disease sampling and monitoring. This project was funded by the New York Wine and Grape Foundation, the Dyson Fund and a USDA Specialty Crop Block Grant administered through the New York Department of Agriculture & Markets.
McFadden-Smith, W., and W.D. Gubler. 2015. "Sour Rot." Compendium of Grape Diseases, Disorders and Pests, Second Edition W.F. Wilcox, W.D. Gubler and J.K. Uyemoto, eds.
Please Note: Illustration(s) are not available due to copyright restrictions.
Caption: The 2017 growing season was difficult for Zinfandel growers battling summer bunch rot in Lodi, Calif.
Caption: This Riesling cluster affected with sour rot is shown shortly before harvest. Note the association of multiple Drosophila fruit flies (arrows) with diseased berries and the lack of fungal mold growth.
Caption: SOUR ROT CONTROL FIELD TRIAL
Caption: EFFECT OF TRAINING SYSTEM ON SOUR ROT DEVELOPMENT
CONTROL OF SOUR ROT WITH ANTI-MICROBIAL SPRAYS INSECTICIDE Mean percent disease control over the untreated check in each year and the mean percent control across all years in which that treatment was administered Antimicrobial treatment, rate and timing (a) Insecticide (b) 2013 2015 2016 None No 0% 0% 0% Yes 9% 57% 40% KMS, 0.5% pre-symptom No 11% Yes 30% KMS, 1 % pre-symptom No 12% 0% 11% Yes 52% 76% 66% KMS, 1 % post-symptom No 8% 34% Yes 46% 46% Kocide 3000, 1.75 pounds/ No 10% acre pre-symptom Yes 54% Fracture, No 14% 32% 32 fluid ounces per Yes 73% 52% acre pre-symptom Fracture, 32 No 12% fluid ounces per Yes 58% acre at 15[degrees] Brix Fracture, No 32% 15% 32 fluid ounces Yes 56% 44% per acre post-symptom Oxidate 2.0, No 31% 3% 1% pre-symptom Yes 84% 55% Oxidate 2.0, No 36% 0% 1% post-symptom Yes 40% 53% Oxidate 2.0, No 27% 1% weekly following Yes 33% first rain after 15[degrees] Brix Oxidate 2.0,1 % weekly following increase in No 47% maximum daily dew point over three consecutive days Yes 33% Antimicrobial treatment, rate and timing (a) Mean None 0% 35% KMS, 0.5% pre-symptom 11% 30% KMS, 1 % pre-symptom 8% 65% KMS, 1 % post-symptom 21% 46% Kocide 3000, 1.75 pounds/ 10% acre pre-symptom 54% Fracture, 23% 32 fluid ounces per 63% acre pre-symptom Fracture, 32 12% fluid ounces per 58% acre at 15[degrees] Brix Fracture, 23% 32 fluid ounces 50% per acre post-symptom Oxidate 2.0, 17% 1% pre-symptom 69% Oxidate 2.0, 18% 1% post-symptom 47% Oxidate 2.0, 27% 1% weekly following 33% first rain after 15[degrees] Brix Oxidate 2.0,1 % weekly following increase in 47% maximum daily dew point over three consecutive days 33% (a) Unless otherwise noted, antimicrobial treatments were applied weekly upon initiation. Pre-symptom treatments started at 15.0[degrees] Brix, and post-symptom treatments after sour rot symptoms were first detected in the vineyard. (b) Insecticide sprays (Delegate [spinetoram] in 2013, Mustang Maxx [zeta-cypermethrin] in 2015 and 2016) were applied to designated rows at label rates weekly beginning at 15.0[degrees] Brix.
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|Title Annotation:||GRAPEGROWING: PRACTICAL WINERY & VINEYARD|
|Comment:||Fruit Flies Play Role in Sour Rot Complex: Training system can impede development of disease.(GRAPEGROWING: PRACTICAL WINERY & VINEYARD)|
|Author:||Hall, Megan; Loeb, Gregory; Wilcox, Wayne|
|Publication:||Wines & Vines|
|Date:||Jan 1, 2018|
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