The heat event that occurred in many parts of Australia in 2009 was the worst on record for the past decade, with air temperatures exceeding 40°C for 14 days. Our aim was to assess the impacts of this heat event on vine performance, including ripening, yield, and gas exchange of Vitis vinifera cv. Semillon grown in a Riverina vineyard. To assess the affect of high temperatures on Semillon grapevines, the vines were covered with a protective layer to reduce radiant heating and were compared with vines exposed to ambient conditions. The heat event had major effects on ripening; reducing the rate of ripening by 50% and delaying harvest ripeness and causing a high incidence of berry shrivel and sunburn. Yield was not affected. Photosynthesis was reduced 35% by the heat event while transpiration increased nearly threefold and was accounted for by increased stomatal conductance. The conclusion of this study was that heat events delayed ripening in Semillon berries and caused a significant reduction in berry quality. Strategies to minimize the radiant load during heat events are required and this study has confirmed a protective layer can reduce canopy temperatures and enhance berry quality.
photosynthesis; rate of ripening; soluble solids concentration; stomatal conductance; transpiration; yield
Shade cloth can be used to protect grapevines from high temperatures. However, the resulting low light intensity is shown to reduce photosynthesis, leading to lower carbon allocation to vegetative growth and sugar accumulation. Protection from heat by shading is, therefore, costly for the carbon economy of the vines.
Background and aims
Covering whole vines with shade cloth is used to protect the vines from heat stress, but may have costs on vine productivity through reduced light availability. Our aim was to assess the carbon balance of vines growing with and without shade to quantify the impact of the covering.
Whole vines were covered with 70 % shade cloth, and shoot leaf area and leaf, stem and bunch growth were followed over two growing seasons. Photosynthesis was measured in situ in all leaves along selected shoots over the growing season. A carbon balance was constructed from the difference in acquisition of carbon and the sequestration of carbon as biomass across the growing seasons.
Shade covering had no initial impact on shoot growth but later reduced leaf growth and later still bunch growth. Stem growth was unaffected. Photosynthetic properties were characteristic of shade leaves, with lower rates and lower light saturation compared with well-exposed leaves. Overall, net photosynthesis was reduced by 40 % by the shade covering and was attributed to the reduced photon flux densities. From the carbon balance, vines were reliant on carbon reserves over 6 weeks after budbreak until current photosynthate increased sufficiently to supply the growth. Shade covering impacted most on biomass accumulation to leaves and bunches at the stage when the vines became autotrophic, consistent with the reduction in carbon acquisition. The markedly high carbon demand by bunches caused a mid-season negative carbon balance, implying that shoots had to draw further on reserves to supply the carbon.
Shade covering over whole grapevines exacerbated the imbalance between the supply of and demand for carbon and greatly reduced vine biomass, especially reproductive allocation. Covering vines with shade cloth to protect the vines from heat events, therefore, had major costs in the carbon economy.
Background and Aims
Daytime root-zone temperature may be a significant factor regulating water flux through plants. Water flux can also occur during the night but nocturnal stomatal response to environmental drivers such as root-zone temperature remains largely unknown.
Here nocturnal and daytime leaf gas exchange was quantified in ‘Shiraz’ grapevines (Vitis vinifera) exposed to three root-zone temperatures from budburst to fruit-set, for a total of 8 weeks in spring.
Despite lower stomatal density, night-time stomatal conductance and transpiration rates were greater for plants grown in warm root-zones. Elevated root-zone temperature resulted in higher daytime stomatal conductance, transpiration and net assimilation rates across a range of leaf-to-air vapour pressure deficits, air temperatures and light levels. Intrinsic water-use efficiency was, however, lowest in those plants with warm root-zones. CO2 response curves of foliar gas exchange indicated that the maximum rate of electron transport and the maximum rate of Rubisco activity did not differ between the root-zone treatments, and therefore it was likely that the lower photosynthesis in cool root-zones was predominantly the result of a stomatal limitation. One week after discontinuation of the temperature treatments, gas exchange was similar between the plants, indicating a reversible physiological response to soil temperature.
In this anisohydric grapevine variety both night-time and daytime stomatal conductance were responsive to root-zone temperature. Because nocturnal transpiration has implications for overall plant water status, predictive climate change models using stomatal conductance will need to factor in this root-zone variable.
Water-use efficiency; leaf gas exchange; CO2 assimilation; grapevine; Vitis vinifera; stomatal conductance; root-zone temperature; ‘Shiraz’; grapevine; soil temperature
Background and Aims
The influence of temperature on the timing of budbreak in woody perennials is well known, but its effect on subsequent shoot growth and architecture has received little attention because it is understood that growth is determined by current temperature. Seasonal shoot development of grapevines (Vitis vinifera) was evaluated following differences in temperature near budbreak while minimizing the effects of other microclimatic variables.
Dormant buds and emerging shoots of field-grown grapevines were heated above or cooled below the temperature of ambient buds from before budbreak until individual flowers were visible on inflorescences, at which stage the shoots had four to eight unfolded leaves. Multiple treatments were imposed randomly on individual plants and replicated across plants. Shoot growth and development were monitored during two growing seasons.
Higher bud temperatures advanced the date of budbreak and accelerated shoot growth and leaf area development. Differences were due to higher rates of shoot elongation, leaf appearance, leaf-area expansion and axillary-bud outgrowth. Although shoots arising from heated buds grew most vigorously, apical dominance in these shoots was reduced, as their axillary buds broke earlier and gave rise to more vigorous lateral shoots. In contrast, axillary-bud outgrowth was minimal on the slow-growing shoots emerging from buds cooled below ambient. Variation in shoot development persisted or increased during the growing season, well after temperature treatments were terminated and despite an imposed soil water deficit.
The data indicate that bud-level differences in budbreak temperature may lead to marked differences in shoot growth, shoot architecture and leaf-area development that are maintained or amplified during the growing season. Although growth rates commonly are understood to reflect current temperatures, these results demonstrate a persistent effect of early-season temperatures, which should be considered in future growth models.
Apical dominance; budbreak; grapevine; growth; leaf expansion; shoot elongation; temperature; Vitis vinifera
Background and Aims
Many physiological processes such as photosynthesis, respiration and transpiration can be strongly influenced by the diurnal patterns of within-tree water potential. Despite numerous experiments showing the effect of water potential on fruit-tree development and growth, there are very few models combining carbohydrate allocation with water transport. The aim of this work was to include a xylem circuit into the functional–structural L-PEACH model.
The xylem modelling was based on an electrical circuit analogy and the Hagen–Poisseuille law for hydraulic conductance. Sub-models for leaf transpiration, soil water potential and the soil–plant interface were also incorporated to provide the driving force and pathway for water flow. The model was assessed by comparing model outputs to field measurements and published knowledge.
The model was able to simulate both the water uptake over a season and the effect of different irrigation treatments on tree development, growth and fruit yield.
This work opens the way to a new field of modelling where complex interactions between water transport, carbohydrate allocation and physiological functions can be simulated at the organ level and describe functioning and behaviour at the tree scale.
Carbon allocation; water stress; xylem; functional–structural plant modelling; plant growth simulation; L-PEACH
Background and Aims
Leaf responses to environmental conditions have been frequently described in fruit trees, but differences among cultivars have received little attention. This study shows that parameters of Farquhar's photosynthesis and Jarvis' stomatal conductance models differed between two apple cultivars, and examines the consequences of these differences for leaf water use efficiency.
Leaf stomatal conductance (gsw), net CO2 assimilation rate (An), respiration (Rd) and transpiration (E) were measured during summer in 8-year-old ‘Braeburn’ and ‘Fuji’ apple trees under well-watered field conditions. Parameters of Farquhar's and Jarvis' models were estimated, evaluated and then compared between cultivars. Leaf carbon isotope discrimination (Δ13C) was measured at the end of the growing season.
A single positive relationship was established between VCmax (maximum carboxylation rate) and Na (leaf nitrogen concentration per unit area), and between Jmax (maximum light-driven electron transport rate) and Na. A higher leaf Rd was observed in ‘Fuji’. The gsw responded similarly to increasing irradiance and leaf temperature in both cultivars. gsw responded to lower vapour pressure deficit in ‘Fuji’ than in ‘Braeburn’. Maximal conductance (gswmax) was significantly smaller and An was more limited by gsw in ‘Braeburn’ than ‘Fuji’. Lower gsw, E and higher intrinsic water use efficiency were shown in ‘Braeburn’ and confirmed by smaller leaf Δ13C compared with ‘Fuji’ leaves.
The use of functional model parameters allowed comparison of the two cultivars and provided evidence of different water use ‘strategies’: ‘Braeburn’ was more conservative in water use than ‘Fuji’, due to stomatal limitation of An, higher intrinsic water use efficiency and lower Δ13C. These physiological traits need to be considered in relation to climate adaptation, breeding of new cultivars and horticultural practice.
Apple; carbon isotope discrimination; leaf nitrogen; leaf temperature; irradiance; Malus × domestica; modelling; photosynthesis; stomata; transpiration; vapour pressure deficit; water use efficiency
Grapevine phylloxera, Daktulosphaira vitifoliae (Fitch) (Hemiptera: Phylloxeridae) is a worldwide pest of Vitis species. It has forms that feed on leaves and roots. Root forms predominate on Vitis vinifera (L.) cultivars, while leaf forms predominate on Vitis species from its native American range. Recently, high densities of D. vitifoliae infestations in leaves of V. vinifera in Brazil, Peru, and Uruguay have been reported. The aims of this study were to determine the seasonal development of grape phylloxera, quantify infestation levels on V. vinifera leaves, and compare them with infestation levels on leaves of a rootstock of American origin. Studies were conducted in two vineyards in Uruguay from 2004–2007. Terminal shoots of 3309 C and Cabernet Sauvignon, Chardonnay, Tannat, Viognier, grafted onto resistant rootstock, were sampled weekly and leaves examined for gall presence and insect life stage. First galls were detected in early October; eggs began to appear within two weeks. Two oviposition peaks occurred by the end of December, and they coincided with bursts of shoot growth. On 3309C rootstock, oviposition peaks were more frequent than on the European cultivars. Based on thermal accumulation, D. vitifoliae could complete eight generations a year in Uruguay. Rootstock 3309C suffered the greatest damage but in some cases was similar to the European cultivars. Damage to Chardonnay, Cabernet Sauvignon and Viognier were also high. There were no galls on Tannat. The 2005–2006 season was characterized by low infestation rates caused by a prolonged drought that affected vegetative growth. There were also differences between vineyards, where the vigorous plants suffering more damage. Leaf galling phylloxera incidence and damage were mainly associated to the cultivar but plant vigor and environmental factors also contributed to increase the incidence.
Daktulosphaira vitifoliae; Hemiptera: Phylloxeridae; Seasonal development; Foliage infestation
A spatially explicit mechanistic model, MAESTRA, was used to separate key parameters affecting transpiration to provide insights into the most influential parameters for accurate predictions of within-crown and within-canopy transpiration. Once validated among Acer rubrum L. genotypes, model responses to different parameterization scenarios were scaled up to stand transpiration (expressed per unit leaf area) to assess how transpiration might be affected by the spatial distribution of foliage properties. For example, when physiological differences were accounted for, differences in leaf width among A. rubrum L. genotypes resulted in a 25% difference in transpiration. An in silico within-canopy sensitivity analysis was conducted over the range of genotype parameter variation observed and under different climate forcing conditions. The analysis revealed that seven of 16 leaf traits had a ≥5% impact on transpiration predictions. Under sparse foliage conditions, comparisons of the present findings with previous studies were in agreement that parameters such as the maximum Rubisco-limited rate of photosynthesis can explain ∼20% of the variability in predicted transpiration. However, the spatial analysis shows how such parameters can decrease or change in importance below the uppermost canopy layer. Alternatively, model sensitivity to leaf width and minimum stomatal conductance was continuous along a vertical canopy depth profile. Foremost, transpiration sensitivity to an observed range of morphological and physiological parameters is examined and the spatial sensitivity of transpiration model predictions to vertical variations in microclimate and foliage density is identified to reduce the uncertainty of current transpiration predictions.
Boundary layer conductance; leaf width; modelling; sensitivity analysis; stomatal conductance; transpiration; water vapour transfer; wind
The aim of this research was to implement a methodology through the generation of a supervised classifier based on the Mahalanobis distance to characterize the grapevine canopy and assess leaf area and yield using RGB images. The method automatically processes sets of images, and calculates the areas (number of pixels) corresponding to seven different classes (Grapes, Wood, Background, and four classes of Leaf, of increasing leaf age). Each one is initialized by the user, who selects a set of representative pixels for every class in order to induce the clustering around them. The proposed methodology was evaluated with 70 grapevine (V. vinifera L. cv. Tempranillo) images, acquired in a commercial vineyard located in La Rioja (Spain), after several defoliation and de-fruiting events on 10 vines, with a conventional RGB camera and no artificial illumination. The segmentation results showed a performance of 92% for leaves and 98% for clusters, and allowed to assess the grapevine’s leaf area and yield with R2 values of 0.81 (p < 0.001) and 0.73 (p = 0.002), respectively. This methodology, which operates with a simple image acquisition setup and guarantees the right number and kind of pixel classes, has shown to be suitable and robust enough to provide valuable information for vineyard management.
clustering; Mahalanobis; Vitis vinifera L.; vineyard; yield assessment
Background and Aims
Plant architecture and its interaction with agronomic practices and environmental constraints are determinants of the structure of the canopy, which is involved in carbon acquisition and fruit quality development. A framework for the quantitative analysis of grapevine (Vitis vinifera) shoot architecture, based on a set of topological and geometrical parameters, was developed for the identification of differences between cultivars and the origins of phenotypic variability.
Two commercial cultivars (‘Grenache N’, ‘Syrah’) with different shoot architectures were grown in pots, in well-irrigated conditions. Shoot topology was analysed, using a hidden semi-Markov chain and variable-order Markov chains to identify deviations from the normal pattern of succession of phytomer types (P0–P1–P2), together with kinematic analysis of shoot axis development. Shoot geometry was characterized by final internode and individual leaf area measurements.
Shoot architecture differed significantly between cultivars. Secondary leaf area and axis length were greater for ‘Syrah’. Secondary leaf area distribution along the main axis also differed between cultivars, with secondary leaves preferentially located towards the basal part of the shoot in ‘Syrah’. The main factors leading to differences in leaf area between the cultivars were: (a) slight differences in main shoot structure, with the supplementary P0 phytomer on the lower part of the shoot in ‘Grenache N’, which bears a short branch; and (b) an higher rate and duration of development of branches bearing by P1–P2 phytomers related to P0 ones at the bottom of the shoot in ‘Syrah’. Differences in axis length were accounted for principally by differences in individual internode morphology, with ‘Syrah’ having significantly longer internodes. This trait, together with a smaller shoot diameter, may account for the characteristic ‘droopy’ habit of ‘Syrah’ shoots.
This study highlights the architectural parameters involved in the phenotypic variability of shoot architecture in two grapevine cultivars. Differences in primary shoot structure and in branch development potential accounted for the main differences in leaf area distribution between the two cultivars. By contrast, shoot shape seemed to be controlled by differences in axis length due principally to differences in internode length.
Architecture; shoot; organogenesis; morphogenesis; branching; leaf area; genotypic variability; Vitis vinifera
Reduction of hydraulic conductance to the canopy has been shown to result in stomatal responses to limit transpiration. To test for similar responses to perturbations of the hydraulic network in leaves, we simultaneously measured leaf gas exchange with spatially explicit chlorophyll-a fluorescence and leaf temperature to examine the effects of cutting a primary leaf vein in Helianthus annuus. We repeated the leaf treatment at each of three different vapor pressure deficits and monitored the short-term dynamics of gas exchange following the treatment. Immediately after treatment, photosynthesis and stomatal conductance (gs) showed a transient “wrong way” response in which photosynthesis declined despite increased gs. Comparisons of fluorescence and temperature across the leaf showed that both photosynthesis and gs were transiently patchy across the measured leaf area, but that the patchiness of the two processes did not correspond in space or time. This suggests that photosynthesis and gs respond to vein cutting-induced cavitation via different mechanisms. Because the stomatal response varied by vapor pressure difference condition but photosynthesis did not, it is likely that gs, but not photosynthesis, responded to a hydraulic signal. In contrast, we hypothesize that photosynthesis declined due to a wound-induced electrical signal that has recently been shown to transiently decrease mesophyll conductance to CO2. The interaction of epidermal hydraulics and the electrical signal across the leaf likely created a patchy pattern of chlorophyll fluorescence and leaf temperature that cannot be explained through the action of a single signal.
stomatal conductance; leaf hydraulic conductance; mesophyll conductance; stomatal patchiness; chlorophyll fluorescence imaging; photosynthesis; transpiration; cavitation
Models seldom consider the effect of leaf-level biochemical acclimation to temperature when scaling forest water use. Therefore, the dependence of transpiration on temperature acclimation was investigated at the within-crown scale in climatically contrasting genotypes of Acer rubrum L., cv. October Glory (OG) and Summer Red (SR). The effects of temperature acclimation on intracanopy gradients in transpiration over a range of realistic forest growth temperatures were also assessed by simulation. Physiological parameters were applied, with or without adjustment for temperature acclimation, to account for transpiration responses to growth temperature. Both types of parameterization were scaled up to stand transpiration (expressed per unit leaf area) with an individual tree model (MAESTRA) to assess how transpiration might be affected by spatial and temporal distributions of foliage properties. The MAESTRA model performed well, but its reproducibility was dependent on physiological parameters acclimated to daytime temperature. Concordance correlation coefficients between measured and predicted transpiration were higher (0.95 and 0.98 versus 0.87 and 0.96) when model parameters reflected acclimated growth temperature. In response to temperature increases, the southern genotype (SR) transpiration responded more than the northern (OG). Conditions of elevated long-term temperature acclimation further separate their transpiration differences. Results demonstrate the importance of accounting for leaf-level physiological adjustments that are sensitive to microclimate changes and the use of provenance-, ecotype-, and/or genotype-specific parameter sets, two components likely to improve the accuracy of site-level and ecosystem-level estimates of transpiration flux.
Intraspecific acclimation; MAESTRA; microclimate; modelling; red maple; temperature acclimation; transpiration
Background and Aims
The bacterium Xylella fastidiosa (Xf), responsible for Pierce's disease (PD) of grapevine, colonizes the xylem conduits of vines, ultimately killing the plant. However, Vitis vinifera grapevine varieties differ in their susceptibility to Xf and numerous other plant species tolerate Xf populations without showing symptoms. The aim of this study was to examine the xylem structure of grapevines with different susceptibilities to Xf infection, as well as the xylem structure of non-grape plant species that support or limit movement of Xf to determine if anatomical differences might explain some of the differences in susceptibility to Xf.
Air and paint were introduced into leaves and stems to examine the connectivity between stem and leaves and the length distribution of their vessels. Leaf petiole and stem anatomies were studied to determine the basis for the free or restricted movement of Xf into the plant.
There were no obvious differences in stem or petiole vascular anatomy among the grape varieties examined, nor among the other plant species that would explain differences in resistance to Xf. Among grape varieties, the more tolerant ‘Sylvaner’ had smaller stem vessel diameters and 20 % more parenchyma rays than the other three varieties. Alternative hosts supporting Xf movement had slightly longer open xylem conduits within leaves, and more connection between stem and leaves, when compared with alternative hosts that limit Xf movement.
Stem–leaf connectivity via open xylem conduits and vessel length is not responsible for differences in PD tolerance among grape varieties, or for limiting bacterial movement in the tolerant plant species. However, it was found that tolerant host plants had narrower vessels and more parenchyma rays, possibly restricting bacterial movement at the level of the vessels. The implications of xylem structure and connectivity for the means and regulation of bacterial movement are discussed.
Grape; grapevine; Vitis vinifera; host; leaf; stem; xylem; Pierce's disease; Xylella fastidiosa
Oryza meridionalis Ng. is a wild relative of Oryza sativa L. found throughout northern Australia where temperatures regularly exceed 35 °C in the monsoon growing season. Heat tolerance in O. meridionalis was established by comparing leaf elongation and photosynthetic rates at 45 °C with plants maintained at 27 °C. By comparison with O. sativa ssp. japonica cv. Amaroo, O. meridionalis was heat tolerant. Elongation rates of the third leaf of O. meridionalis declined by 47% over 24 h at 45 °C compared with a 91% decrease for O. sativa. Net photosynthesis was significantly higher in O. sativa at 27 °C whereas the two species had the same assimilation rates at 45 °C. The leaf proteome and expression levels of individual heat-responsive genes provided insight into the heat response of O. meridionalis. After 24 h of heat exposure, many enzymes involved in the Calvin Cycle were more abundant, while mRNA of their genes generally decreased. Ferredoxin-NADP(H) oxidoreductase, a key enzyme in photosynthetic electron transport had both reduced abundance and gene expression, suggesting light reactions were highly susceptible to heat stress. Rubisco activase was strongly up-regulated after 24 h of heat, with the large isoform having the largest relative increase in protein abundance and a significant increase in gene expression. The protective proteins Cpn60, Hsp90, and Hsp70 all increased in both protein abundance and gene expression. A thiamine biosynthesis protein (THI1), previously shown to act protectively against stress, increased in abundance during heat, even as thiamine levels fell in O. meridionalis.
Calvin Cycle; dark reaction; ferredoxin-NADP(H) oxidoreductase; heat shock protein; heat stress; leaf elongation; O. meridionalis; Rubisco activase; thiamine biosynthesis protein (THI1)
To understand the physiological basis of genetic variation and resulting quantitative trait loci (QTLs) for photosynthesis in a rice (Oryza sativa L.) introgression line population, 13 lines were studied under drought and well-watered conditions, at flowering and grain filling. Simultaneous gas exchange and chlorophyll fluorescence measurements were conducted at various levels of incident irradiance and ambient CO2 to estimate parameters of a model that dissects photosynthesis into stomatal conductance (g
s), mesophyll conductance (g
m), electron transport capacity (J
max), and Rubisco carboxylation capacity (V
cmax). Significant genetic variation in these parameters was found, although drought and leaf age accounted for larger proportions of the total variation. Genetic variation in light-saturated photosynthesis and transpiration efficiency (TE) were mainly associated with variation in g
s and g
m. One previously mapped major QTL of photosynthesis was associated with variation in g
s and g
m, but also in J
max and V
cmax at flowering. Thus, g
s and g
m, which were demonstrated in the literature to be responsible for environmental variation in photosynthesis, were found also to be associated with genetic variation in photosynthesis. Furthermore, relationships between these parameters and leaf nitrogen or dry matter per unit area, which were previously found across environmental treatments, were shown to be valid for variation across genotypes. Finally, the extent to which photosynthesis rate and TE can be improved was evaluated. Virtual ideotypes were estimated to have 17.0% higher photosynthesis and 25.1% higher TE compared with the best genotype investigated. This analysis using introgression lines highlights possibilities of improving both photosynthesis and TE within the same genetic background.
drought; genetic variation; mesophyll conductance; modelling; Oryza sativa L.; photosynthesis; rice; stomatal conductance
Background and Aims
Nitrogen (N) is a major factor affecting yield gain of crops under elevated atmospheric carbon dioxide concentrations [CO2]. It is well established that elevated [CO2] increases root mass, but there are inconsistent reports on the effects on N uptake capacity per root mass. In the present study, it was hypothesized that the responses of N uptake capacity would change with the duration of exposure to elevated [CO2].
The hypothesis was tested by measuring N uptake capacity in rice plants exposed to long-term and short-term [CO2] treatments at different growth stages in plants grown under non-limiting N conditions in hydroponic culture. Seasonal changes in photosynthesis rate and transpiration rate were also measured.
In the long-term [CO2] study, leaf photosynthetic responses to intercellular CO2 concentration (Ci) were not affected by elevated [CO2] before the heading stage, but the initial slope in this response was decreased by elevated [CO2] at the grain-filling stage. Nitrate and ammonium uptake capacities per root dry weight were not affected by elevated [CO2] at panicle initiation, but thereafter they were reduced by elevated [CO2] by 31–41 % at the full heading and mid-ripening growth stages. In the short-term study (24 h exposures), elevated [CO2] enhanced nitrate and ammonium uptake capacities at the early vegetative growth stage, but elevated [CO2] decreased the uptake capacities at the mid-reproductive stage.
This study showed that N uptake capacity was downregulated under long-term exposure to elevated [CO2] and its response to elevated [CO2] varied greatly with growth stage.
Acclimation; elevated CO2; nitrogen uptake capacity; Oryza sativa; photosynthesis rate; rice; root development; transpiration rate
Herbivory reduces leaf area, disrupts the function of leaves, and ultimately alters yield and productivity. Herbivore damage to foliage typically is assessed in the field by measuring the amount of leaf tissue removed and disrupted. This approach assumes the remaining tissues are unaltered, and plant photosynthesis and water balance function normally. However, recent application of thermal and fluorescent imaging technologies revealed that alterations to photosynthesis and transpiration propagate into remaining undamaged leaf tissue.
Scope and Conclusions
This review briefly examines the indirect effects of herbivory on photosynthesis, measured by gas exchange or chlorophyll fluorescence, and identifies four mechanisms contributing to the indirect suppression of photosynthesis in remaining leaf tissues: severed vasculature, altered sink demand, defence-induced autotoxicity, and defence-induced down-regulation of photosynthesis. We review the chlorophyll fluorescence and thermal imaging techniques used to gather layers of spatial data and discuss methods for compiling these layers to achieve greater insight into mechanisms contributing to the indirect suppression of photosynthesis. We also elaborate on a few herbivore-induced gene-regulating mechanisms which modulate photosynthesis and discuss the difficult nature of measuring spatial heterogeneity when combining fluorescence imaging and gas exchange technology. Although few studies have characterized herbivore-induced indirect effects on photosynthesis at the leaf level, an emerging literature suggests that the loss of photosynthetic capacity following herbivory may be greater than direct loss of photosynthetic tissues. Depending on the damage guild, ignoring the indirect suppression of photosynthesis by arthropods and other organisms may lead to an underestimate of their physiological and ecological impacts.
Chlorophyll fluorescence imaging; thermography; plant–insect interactions; spatial patterns; autotoxicity; induced defences; jasmonates
Background and Aims
Grapevine (Vitis spp.) cold hardiness varies dynamically throughout the dormant season, primarily in response to changes in temperature. The development and possible uses of a discrete-dynamic model of bud cold hardiness for three Vitis genotypes are described.
Iterative methods were used to optimize and evaluate model parameters by minimizing the root mean square error between observed and predicted bud hardiness, using up to 22 years of low-temperature exotherm data. Three grape cultivars were studied: Cabernet Sauvignon, Chardonnay (both V. vinifera) and Concord (V. labruscana). The model uses time steps of 1 d along with the measured daily mean air temperature to calculate the change in bud hardiness, which is then added to the hardiness from the previous day. Cultivar-dependent thermal time thresholds determine whether buds acclimate (gain hardiness) or deacclimate (lose hardiness).
The parameterized model predicted bud hardiness for Cabernet Sauvignon and Chardonnay with an r2 = 0·89 and for Concord with an r2 = 0·82. Thermal time thresholds and (de-)acclimation rates changed between the early and late dormant season and were cultivar dependent but independent of each other. The timing of these changes was also unique for each cultivar. Concord achieved the greatest mid-winter hardiness but had the highest deacclimation rate, which resulted in rapid loss of hardiness in spring. Cabernet Sauvignon was least hardy, yet maintained its hardiness latest as a result of late transition to eco-dormancy, a high threshold temperature required to induce deacclimation and a low deacclimation rate.
A robust model of grapevine bud cold hardiness was developed that will aid in the anticipation of and response to potential injury from fluctuations in winter temperature and from extreme cold events. The model parameters that produce the best fit also permit insight into dynamic differences in hardiness among genotypes.
Cold hardiness; cold injury; differential thermal analysis; discrete model; grapevine; low-temperature exotherm; Vitis labruscana; Vitis vinifera
• Background and Aims Kaolin applications have been used to mitigate the negative effects of water and heat stress on plant physiology and productivity with variable results, ranging from increased to decreased yields and photosynthetic rates. The mechanisms of action of kaolin applications are not clear: although the increased albedo reduces leaf temperature and the consequent heat stress, it also reduces the light available for photosynthesis, possibly offsetting benefits of lower temperature. The objective of this study was to investigate which of these effects are prevalent and under which conditions.
• Methods A 6 % kaolin suspension was applied on well-irrigated and water-stressed walnut (Juglans regia) and almond (Prunus dulcis) trees. Water status (i.e. stem water potential, Ψs), gas exchange (i.e. light-saturated CO2 assimilation rate, Amax; stomatal conductance, gs), leaf temperature (Tl) and physiological relationships in treated and control trees were then measured and compared.
• Key Results In both species, kaolin did not affect the daily course of Ψs whereas it reduced Amax by 1–4 μmol CO2 m–2 s–1 throughout the day in all combinations of species and irrigation treatments. Kaolin did not reduce gs in any situation. Consequently, intercellular CO2 concentration (Ci) was always greater in treated trees than in controls, suggesting that the reduction of Amax with kaolin was not due to stomatal limitations. Kaolin reduced leaf temperature (Tl) by about 1–3 °C and leaf-to-air vapour pressure difference (VPDl) by about 0·1–0·7 kPa. Amax was lower at all values of gs, Tl and VPDl in kaolin-treated trees. Kaolin affected the photosynthetic response to the photosynthetically active radiation (PAR) in almond leaves: kaolin-coated leaves had similar dark respiration rates and light-saturated photosynthesis, but a higher light compensation point and lower apparent quantum yield, while the photosynthetic light-response curve saturated at higher PAR. When these parameters were used to model the photosynthetic response curve to PAR, it was estimated that the kaolin film allowed 63 % of the incident PAR to reach the leaf.
• Conclusions The main effect of kaolin application was the reduction, albeit minor, of photosynthesis, which appeared to be related to the shading of the leaves. The reduction in Tl and VPDl with kaolin did not suffice to mitigate the adverse effects of heat and water stress on Amax.
Juglans regia; kaolin particle film; photosynthesis; Prunus dulcis; stomatal conductance; water potential; stress
Phytoplasmas are bacteria without cell walls from the class Mollicutes. They are obligate intracellular plant pathogens which cause diseases in hundreds of economically important plants including the grapevine (Vitis vinifera). Knowledge of their biology and the mechanisms of their interactions with hosts is largely unknown because they are uncultivable and experimentally inaccessible in their hosts. We detail here the global transcriptional profiling in grapevine responses to phytoplasmas. The gene expression patterns were followed in leaf midribs of grapevine cv. 'Chardonnay' naturally infected with a phytoplasma from the stolbur group 16SrXII-A, which is associated with the grapevine yellows disease 'Bois noir'.
We established an on field experimental system in a productive vineyard that allowed application of molecular tools in a plant natural environment. Global transcription profiles of infected samples were compared with the healthy ones using microarray datasets and metabolic pathway analysis software (MapMan). The two-year-long experiment revealed that plant genes involved in primary and secondary metabolic pathways were changed in response to infection and that these changes might support phytoplasma nutrition. A hypothesis that phytoplasmas interact with the plant carbohydrate metabolism was proven and some possibilities how the products of this pathway might be utilized by phytoplasmas are discussed. In addition, several photosynthetic genes were largely down-regulated in infected plants, whereas defense genes from the metabolic pathway leading to formation of flavonoids and some PR proteins were significantly induced. Few other genes involved in defense-signaling were differentially expressed in healthy and infected plants. A set of 17 selected genes from several differentially expressed pathways was additionally analyzed with quantitative real-time PCR and confirmed to be suitable for a reliable classification of infected plants and for the characterization of susceptibility features in the field conditions.
This study revealed some fundamental aspects of grapevine interactions with the stolbur 'Bois noir' phytoplasma in particular and some plant interactions with phytoplasmas in general. In addition, the results of the study will likely have an impact on grape improvement by yielding marker genes that can be used in new diagnostic assays for phytoplasmas or by identifying candidate genes that contribute to the improved properties of grape.
Eutypa dieback is a vascular disease that may severely affect vineyards throughout the world. In the present work, microarrays were made in order (i) to improve our knowledge of grapevine (Vitis vinifera cv. Cabernet-Sauvignon) responses to Eutypa lata, the causal agent of Eutypa dieback; and (ii) to identify genes that may prevent symptom development. Qiagen/Operon grapevine microarrays comprising 14 500 probes were used to compare, under three experimental conditions (in vitro, in the greenhouse, and in the vineyard), foliar material of infected symptomatic plants (S+R+), infected asymptomatic plants (S–R+), and healthy plants (S–R–). These plants were characterized by symptom notation after natural (vineyard) or experimental (in vitro and greenhouse) infection, re-isolation of the fungus located in the lignified parts, and the formal identification of E. lata mycelium by PCR. Semi-quantitative real-time PCR experiments were run to confirm the expression of some genes of interest in response to E. lata. Their expression profiles were also studied in response to other grapevine pathogens (Erysiphe necator, Plasmopara viticola, and Botrytis cinerea). (i) Five functional categories of genes, that is those involved in metabolism, defence reactions, interaction with the environment, transport, and transcription, were up-regulated in S+R+ plants compared with S–R– plants. These genes, which cannot prevent infection and symptom development, are not specific since they were also up-regulated after infection by powdery mildew, downy mildew, and black rot. (ii) Most of the genes that may prevent symptom development are associated with the light phase of photosynthesis. This finding is discussed in the context of previous data on the mode of action of eutypin and the polypeptide fraction secreted by Eutypa.
Eutypa dieback; Eutypa lata; grapevine; microarrays; transcriptome; Vitis vinifera
Grapevine roots can be exposed to a range of temperatures at any particular moment because the root system can explore large volumes of soil over great depths and distances. A split-pot experiment was designed to assess how vegetative and reproductive development respond to partial and whole root-zone warming following winter dormancy. Simultaneous cooling and warming of parts of the root system slowed shoot elongation, leaf expansion and berry development compared to plants with a fully warmed root-zone, but not to the same extent as those with a fully cooled root-zone.
Heterogeneity in root-zone temperature both vertically and horizontally may contribute to the uneven vegetative and reproductive growth often observed across vineyards. An experiment was designed to assess whether the warmed half of a grapevine root zone could compensate for the cooled half in terms of vegetative growth and reproductive development. We divided the root system of potted Shiraz grapevines bilaterally and applied either a cool or a warm treatment to each half from budburst to fruit set. Shoot growth and inflorescence development were monitored over the season. Simultaneous cooling and warming of parts of the root system decreased shoot elongation, leaf emergence and leaf expansion below that of plants with a fully warmed root zone, but not to the same extent as those with a fully cooled root zone. Inflorescence rachis length, flower number and berry number after fertilization were smaller only in those vines exposed to fully cooled root zones. After terminating the treatments, berry enlargement and the onset of veraison were slowed in those vines that had been exposed to complete or partial root-zone cooling. Grapevines exposed to partial root-zone cooling were thus delayed in vegetative and reproductive development, but the inhibition was greater in those plants whose entire root system had been cooled.
Berry composition; grapevine; Shiraz; shoot growth; soil temperature; split pot; Vitis vinifera.
Background and Aims
Global climate models predict decreases in leaf stomatal conductance and transpiration due to increases in atmospheric CO2. The consequences of these reductions are increases in soil moisture availability and continental scale run-off at decadal time-scales. Thus, a theory explaining the differential sensitivity of stomata to changing atmospheric CO2 and other environmental conditions must be identified. Here, these responses are investigated using optimality theory applied to stomatal conductance.
An analytical model for stomatal conductance is proposed based on: (a) Fickian mass transfer of CO2 and H2O through stomata; (b) a biochemical photosynthesis model that relates intercellular CO2 to net photosynthesis; and (c) a stomatal model based on optimization for maximizing carbon gains when water losses represent a cost. Comparisons between the optimization-based model and empirical relationships widely used in climate models were made using an extensive gas exchange dataset collected in a maturing pine (Pinus taeda) forest under ambient and enriched atmospheric CO2.
Key Results and Conclusion
In this interpretation, it is proposed that an individual leaf optimally and autonomously regulates stomatal opening on short-term (approx. 10-min time-scale) rather than on daily or longer time-scales. The derived equations are analytical with explicit expressions for conductance, photosynthesis and intercellular CO2, thereby making the approach useful for climate models. Using a gas exchange dataset collected in a pine forest, it is shown that (a) the cost of unit water loss λ (a measure of marginal water-use efficiency) increases with atmospheric CO2; (b) the new formulation correctly predicts the condition under which CO2-enriched atmosphere will cause increasing assimilation and decreasing stomatal conductance.
Economics of gas exchange; free air CO2 enrichment; marginal water-use efficiency; photosynthesis; Pinus taeda; stomatal conductance; stomatal optimization
Background and Aims
The rate of photosynthesis in paddy rice often decreases at noon on sunny days because of water stress, even under submerged conditions. Maintenance of higher rates of photosynthesis during the day might improve both yield and dry matter production in paddy rice. A high-yielding indica variety, ‘Habataki’, maintains a high rate of leaf photosynthesis during the daytime because of the higher hydraulic conductance from roots to leaves than in the standard japonica variety ‘Sasanishiki’. This research was conducted to characterize the trait responsible for the higher hydraulic conductance in ‘Habataki’ and identified a chromosome region for the high hydraulic conductance.
Hydraulic conductance to passive water transport and to osmotic water transport was determined for plants under intense transpiration and for plants without transpiration, respectively. The varietal difference in hydraulic conductance was examined with respect to root surface area and hydraulic conductivity (hydraulic conductance per root surface area, Lp). To identify the chromosome region responsible for higher hydraulic conductance, chromosome segment substitution lines (CSSLs) derived from a cross between ‘Sasanishiki’ and ‘Habataki’ were used.
The significantly higher hydraulic conductance resulted from the larger root surface area not from Lp in ‘Habataki’. A chromosome region associated with the elevated hydraulic conductance was detected between RM3916 and RM2431 on the long arm of chromosome 4. The CSSL, in which this region was substituted with the ‘Habataki’ chromosome segment in the ‘Sasanishiki’ background, had a larger root mass than ‘Sasanishiki’.
The trait for increasing plant hydraulic conductance and, therefore, maintaining the higher rate of leaf photosynthesis under the conditions of intense transpiration in ‘Habataki’ was identified, and it was estimated that there is at least one chromosome region for the trait located on chromosome 4.
Chromosome segment substitution lines; diffusive conductance; hydraulic conductance; photosynthetic rate; quantitative trait locus; rice; Oryza sativa; root hydraulic conductivity
Grapevine flower development and fruit set are influenced by cold nights in the vineyard. To investigate the impact of cold stress on carbon metabolism in the inflorescence, we exposed the inflorescences of fruiting cuttings to chilling and freezing temperatures overnight and measured fluctuations in photosynthesis and sugar content. Whatever the temperature, after the stress treatment photosynthesis was modified in the inflorescence, but the nature of the alteration depended on the intensity of the cold stress. At 4°C, photosynthesis in the inflorescence was impaired through non-stomatal limitations, whereas at 0°C it was affected through stomatal limitations. A freezing night (−3°C) severely deregulated photosynthesis in the inflorescence, acting primarily on photosystem II. Cold nights also induced accumulation of sugars. Soluble carbohydrates increased in inflorescences exposed to −3°C, 0°C and 4°C, but starch accumulated only in inflorescences of plants treated at 0 and −3°C. These results suggest that inflorescences are able to cope with cold temperatures by adapting their carbohydrate metabolism using mechanisms that are differentially induced according to stress intensity.