El fenómeno de la cavitación en vid… : descifrando los mecanismos implicados

Cavitation is a physiological dysfunction that takes place in the xylem of water stressed plants. It leads to a loss of hydraulic conductance (kL) as the vessels are filled with air. This impacts water supply, water potential (ΨL) and canopy hydration. Stomatal clossure is an effective response upon diminishing momentary or seasonal foliar hydraulic contents. Depending on each type of plant, stomata may close preventing catastrophic cavitations. This research intended to understand how stomatal control acts upon cavitation events in two contrasting grapevine varieties, Syrah and Grenache. A mechanistic was developed model based on the water and vapour fluxes, kL, stomata conductance (gs), and the vulnerability to cavitation of the xylematic tissue. The theoretical model explains how plants respond to drought and avoid catastrophic cavitation. Water stressed grapevines couple their gs with their kL in order to avoid embolism. It is not stomatal closure, by istself, the controlling mechanism. Grapevines under mild water stress, do not need to completely close their stomata in order to avoid cavitation, therefore, photosynthesis is not completely impeded, and the cost in terms of carbon assimilation is less than expected for other species.La cavitación es una disfunción fisiológica que ocurre en el xilema de las plantas bajo déficit hídrico, y que entraña una pérdida de su conductancia hidráulica (kL), cuando algunos vasos se llenan de aire. Esto incide negativamente sobre la oferta de agua y afecta el potencial hídrico foliar (ΨL) y la hidratación de la canopia. El cierre estomático es una respuesta efectiva ante la disminución del contenido hídrico. Dependiendo de la especie vegetal, los estomas suelen cerrase para evitar la cavitación catastrófica. Mediante un modelo mecanístico, que se construyó teniendo en cuenta los flujos de agua y vapor, las kL y conductancia estomática (gs), y la vulnerabilidad del xilema a cavitar; se probó que gs no es la única variable responsable de frenar la embolia. Se determinó que gs y kL están íntimamente asociadas y que este acople entre ambas conductancias es lo que frena la embolia. Se concluyó que, en la vid y bajo niveles de estrés hídrico moderado, no es necesario un cierre estomático para controlar la cavitación. Por esto, el mecanismo de control de la cavitación en la vid no conlleva un costo en términos de intercambio gaseoso.Fil: Hugalde, Inés. Instituto Nacional de Tecnología Agropecuaria (Argentina). Estación Experimental Agropecuaria Mendoza.Fil: Bonada, Marcos. South Australian Research and Development InstituteFil: Vila, Hernán. Instituto Nacional de Tecnología Agropecuaria (Argentina). Estación Experimental Agropecuaria Mendoza

Relaciones entre cavitación y cierre estomático en vides sometidas a déficit hídrico

La cavitación es una disfunción fisiológica que ocurre en el xilema de las plantas cuando estas están bajo déficit hídrico, y que entraña una pérdida de su conductancia hidráulica (kL), cuando algunos vasos se llenan de aire. Esto incide negativamente sobre la oferta de agua y afecta el potencial hídrico foliar (ΨL) y la hidratación de la canopia. El cierre estomático es una respuesta efectiva ante la disminución del contenido hídrico. Dependiendo de la especie vegetal, los estomas suelen cerrase para evitar la cavitación catastrófica. Una planta poco vulnerable a cavitar puede mantenerlos abiertos por más tiempo y por lo tanto seguir fotosintetizando. Por el contrario, plantas vulnerables a la cavitación deben cerrar sus estomas con anterioridad para poder evitar cavitaciones graves. En este trabajo se estudió, el ajuste estomático como mecanismo para evitar la cavitación en cuatro variedades contrastantes de vid (Grenache, Syrah, Malbec y Chardonnay). Se hipotetizó que las vides sometidas a déficit hídrico disminuyen gs para evitar sufrir cavitaciones catastróficas y que algunas variedades de vid, cuando crecen bajo restricción hídrica, se aclimatan mejor desarrollando un ajuste estomático más preciso, ó un sistema conductor menos vulnerable a cavitar, o ambas. Se diseñó un experimento aleatorizado en invernáculo donde se probaron dos situaciones hídricas edáficas (100% y 50% de FTSW). Luego de dos meses, se midieron, a lo largo del día, las variables gs, transpiración y potencial hídrico. Luego se construyeron las curvas de cavitación y se calculó la conduactancia hidráulica de la planta (kL) y el embolismo a lo largo del día. Finalmente se obtuvo la relación entre gs, kL y el embolismo. Mediante un modelo mecanístico, que se construyó teniendo en cuenta los flujos de agua y vapor, las kL y gs, y la vulnerabilidad del xilema a cavitar; se probó que gs no es la única variable responsable de frenar la embolia. Se determinó que gs y kL están íntimamente asociadas y que este acople entre ambas conductancias es lo que frena la embolia. Se concluyó que, en la vid y bajo niveles de estrés hídrico moderado, no es necesario un cierre estomático para controlar la cavitación, sino una disminución de la diferencia entre gs y kL (Δgs). Por esto, el mecanismo de control de la cavitación en la vid no conlleva un costo en términos de intercambio gaseoso. También se descubrió que bajo niveles moderados de déficit hídrico la vulnerabilidad xilemática no aumenta con respecto a las plantas que no sufren déficit, sin importar de qué variedad se trate.Cavitation is a physiological dysfunction that takes place in the xylem of plants under water stress. It leads to a loss of hydraulic conductance as the vessels are filled with air. This has a negative impact on water supply and affects ΨL and canopy hydration. Stomatal clossure is an effective response upon diminishing momentary or seasonal foliar hydraulic contents. Depending on each type of plant, stomata may close preventing catastrophic cavitations. A not vulnerable to cavitation plant may maintain higher stomatal conductances (gs) and therefore greater photosynthesis, by keeping opened stomata for longer periods of time. On the other hand, vulnerable plants should close their stomata in order to prevent runaway embolism and lost of hydraulic conductance. This research intended to understand how stomatal control acts upon cavitation events in four contrasting grapevine varieties (Grenache, Syral, Malbec and Chardonnay). We hypothesized that water stressed grapevines, reduce gs in order to avoid catastrophic embolism, and that some varieties, when grown under water stress, may acclimatize themselves by the development of a precise stomatal control, or a less vulnerable xylem, or both. A randomized experimental plot inside a greenhouse was conducted (two FTSW treatmentes were determined, 100% and 50%). Quantitative measurements of gs, transpiration, photosynthesis and stem water potential were assessed from predawn to 4 pm, every one our. Two months later, cavitation curves were constructed, and the level of embolism reached along the day and plant hydraulic conductance, were calculated. Finally the relationship among gs, kL and embolism was determined. By means of a mechanistical model that was constructed based on the water and vapour fluxes, kL, gs, and the vulnerability to cavitation of the xylematic tissue, we probed that gs is not the only variable that controls cavitation. It was determined that gs is coupled with kL, and this coupling is the responsible mechanism that achieves embolism control. This leaded us to conclude that grapevines under mild water stress, do not need to close their stomata in order to avoid cavitation. They only need a midmorning decrease in the difference between gs and kL (Δgs), therefore, no cost in terms of carbon assimilation is required.Fil: Hugalde, Inés Pilar. Universidad Nacional de Cuyo. Facultad de Ciencias Agrarias

Studying Growth and Vigor as Quantitative Traits in Grapevine Populations

Vigor is considered as a propensity to assimilate, store, and/or use nonstructural carbohydrates for producing large canopies, and it is associated with high metabolism and fast growth. Growth involves cell expansion and cell division. Cell division depends on hormonal and metabolic processes. Cell expansion occurs because cell walls are extensible, meaning they deform under the action of tensile forces, generally caused by turgor. There is increasing interest in understanding the genetic basis of vigor and biomass production. It is well established that growth and vigor are quantitative traits and their genetic architecture consists of a big number of genes with small individual effects. The search for groups of genes with small individual effects, which control a specific quantitative trait, is performed by QTL analysis and genetic mapping. Today, several linkage maps are available, like “Syrah” × “grenache,” “Riesling” × “Cabernet Sauvignon,” and “Ramsey” × Vitis riparia. This last progeny segregates for vigor and constituted an interesting tool for our genetic studies on growth

Prioritization of vigor QTL-associated genes for future genome-directed Vitis breeding: Priorización de genes relacionados a vigor en Vitis

Vigor control in grapevine may become especially important under climate change. A better understanding of gene-phenotype relationships is required in order to exploit plant genomics for breeding purposes. This research aims to use quantitative trait loci (QTLs) for vigor identified in the progeny from a cross of Ramsey (Vitis champinii) × Riparia Gloire (V. riparia). Genes located 700 kb up and downstream from each QTL position were interrogated for functional enrichment through ShinyGO online tool, based on the gene ontology annotation of Vitis vinifera PN40024. Key biological processes like phloem and xylem development, cell cycle, response to hormones, amino acid transport, tissue development, sugar metabolism, nitrogen transport, and stress/immune responses, showed functional enrichment. Integral response to light and auxin might be required for fine molecular tuning of vegetative growth in Vitis. Fifty out of 1318 candidate genes were prioritized, reducing their amount to a manageable number of candidates for further directed breeding strategies. Highlights Plant vigor control may become especially important under climate change. Genes from various vigor-related QTLs were interrogated for functional enrichment. The analysis reduced candidate gene number based on marker proximity and functional enrichment, constituting a suitable shortcut for target-directed genome-guided breeding strategies. Three TFs are strong candidates for targeted breeding: TIF - HY5, TIF - SUS1, TIF - VOZ1 potentially enhance growth by relating light response to hormone activation, and then to photosynthesis and morphogenesis.  Vigor control in grapevine may become especially important under climate change. A better understanding of gene-phenotype relationships is required in order to exploit plant genomics for breeding purposes. This research aims to use quantitative trait loci (QTLs) for vigor identified in the progeny from a cross of Ramsey (Vitis champinii) × Riparia Gloire (V. riparia). Genes located 700 kb up and downstream from each QTL position were interrogated for functional enrichment through ShinyGO online tool, based on the gene ontology annotation of Vitis vinifera PN40024. Key biological processes like phloem and xylem development, cell cycle, response to hormones, amino acid transport, tissue development, sugar metabolism, nitrogen transport, and stress/immune responses, showed functional enrichment. Integral response to light and auxin might be required for fine molecular tuning of vegetative growth in Vitis. Fifty out of 1318 candidate genes were prioritized, reducing their amount to a manageable number of candidates for further directed breeding strategies. Highlights Plant vigor control may become especially important under climate change. Genes from various vigor-related QTLs were interrogated for functional enrichment. The analysis reduced candidate gene number based on marker proximity and functional enrichment, constituting a suitable shortcut for target-directed genome-guided breeding strategies. Three TFs are strong candidates for targeted breeding: TIF - HY5, TIF - SUS1, TIF - VOZ1 potentially enhance growth by relating light response to hormone activation, and then to photosynthesis and morphogenesis

Prioritization of vigor QTL-associated genes for future genome-directed Vitis breeding

Vigor control in grapevine may become especially important under climate change. A better understanding of gene-phenotype relationships is required in order to exploit plant genomics for breeding purposes. This research aims to use quantitative trait loci (QTLs) for vigor identified in the progeny from a cross of Ramsey (Vitis champinii) × Riparia Gloire (V. riparia). Genes located 700 kb up and downstream from each QTL position were interrogated for functional enrichment through ShinyGO online tool, based on the gene ontology annotation of Vitis vinifera PN40024. Key biological processes like phloem and xylem development, cell cycle, response to hormones, amino acid transport, tissue development, sugar metabolism, nitrogen transport, and stress/immune responses, showed functional enrichment. Integral response to light and auxin might be required for fine molecular tuning of vegetative growth in Vitis. Fifty out of 1318 candidate genes were prioritized, reducing their amount to a manageable number of candidates for further directed breeding strategies

Modeling vegetative vigour in grapevine: unraveling underlying mechanisms

Mechanistic modeling constitutes a powerful tool to unravel complex biological phenomena. This study describes the construction of a mechanistic, dynamic model for grapevine plant growth and canopy biomass (vigor). To parametrize and validate the model, the progeny from a cross of Ramsey (Vitis champinii) × Riparia Gloire (V. riparia) was evaluated. Plants with different vigor were grown in a greenhouse during the summer of 2014 and 2015. One set of plants was grafted with Cabernet Sauvignon. Shoot growth rate (b), leaf area (LA), dry biomass, whole plant and root specific hydraulic conductance (kH and Lpr), stomatal conductance (gs), and water potential (Ψ) were measured. Partitioning indices and specific leaf area (SLA) were calculated. The model includes an empirical fit of a purported seasonal pattern of bioactive GAs based on published seasonal evolutionary levels and reference values. The model provided a good fit of the experimental data, with R = 0.85. Simulation of single trait variations defined the individual effect of each variable on vigor determination. The model predicts, with acceptable accuracy, the vigor of a young plant through the measurement of Lpr and SLA. The model also permits further understanding of the functional traits that govern vigor, and, ultimately, could be considered useful for growers, breeders and those studying climate change