The Evolution of Mechanisms Driving the Stomatal Response to Vapor Pressure Deficit

Stomatal responses to vapor pressure deficit (VPD) are a principal means by which vascular land plants regulate daytime transpiration. While much work has focused on characterizing and modeling this response, there remains no consensus as to the mechanism that drives it. Explanations range from passive regulation by leaf hydration to biochemical regulation by the phytohormone abscisic acid (ABA). We monitored ABA levels, leaf gas exchange, and water status in a diversity of vascular land plants exposed to a symmetrical, mild transition in VPD. The stomata in basal lineages of vascular plants, including gymnosperms, appeared to respond passively to changes in leaf water status induced by VPD perturbation, with minimal changes in foliar ABA levels and no hysteresis in stomatal action. In contrast, foliar ABA appeared to drive the stomatal response to VPD in our angiosperm samples. Increased foliar ABA level at high VPD in angiosperm species resulted in hysteresis in the recovery of stomatal conductance; this was most pronounced in herbaceous species. Increased levels of ABA in the leaf epidermis were found to originate from sites of synthesis in other parts of the leaf rather than from the guard cells themselves. The transition from a passive regulation to ABA regulation of the stomatal response to VPD in the earliest angiosperms is likely to have had critical implications for the ecological success of this lineage.Plants continuously regulate transpiration by controlling the aperture of the stomatal pores on the surface of the leaf. The principal atmospheric determinant of stomatal aperture is the humidity of the air, which can be expressed as the vapor pressure difference between the leaf and the atmosphere. Stomatal responses to atmospheric vapor pressure deficit (VPD) have been well characterized across the diversity of vascular plant species (Darwin, 1898; Lange et al., 1971; Turner et al., 1984; Franks and Farquhar, 1999; Oren et al., 1999; Brodribb and McAdam, 2011; Mott and Peak, 2013), with stomata typically closing at high VPD and opening at low VPD. This comprehensive characterization has allowed for the development of highly effective empirical and mechanistic models of leaf gas exchange that provide robust predictions of the responses of transpiration to changes in VPD (Buckley et al., 2003; Katul et al., 2009; Damour et al., 2010; Medlyn et al., 2011). Despite the success of this modeling, the mechanism for the stomatal response to VPD remains poorly understood (Damour et al., 2010). Different hypotheses range from one extreme, whereby stomata respond passively through changes in leaf water content induced by the VPD or humidity perturbation (Lange et al., 1971; Mott and Peak, 2013), to the other extreme, whereby stomata close uniquely in response to the phytohormone abscisic acid (ABA; Xie et al., 2006; Bauer et al., 2013).From the earliest recognition that stomata open and close by changes in guard cell turgor (Heath, 1938), there have been many attempts to link the passive changes in water status that occur during VPD or humidity transitions with stomatal responses to VPD or humidity (Lange et al., 1971; Mott and Peak, 2013). Studies have suggested that changes in atmospheric water content passively drive stomatal responses by changing bulk leaf water status, which in turn changes guard cell turgor (Oren et al., 1999), or alternatively by changing guard cell turgor directly (Mott and Peak, 2013). Models based on these entirely passive processes are highly effective in predicting steady-state stomatal conductance (g_s) in response to changes in VPD or humidity in angiosperms (Mott and Peak, 2013).While hydraulic models provide robust predictions of steady-state g_s, they are less effective at predicting the dynamic responses of stomata to short-term perturbations, particularly with respect to the wrong-way responses that typically occur as transients (Buckley, 2005), as well as feed-forward behavior (Farquhar, 1978; Bunce, 1997; Franks et al., 1997; Tardieu and Simonneau, 1998; Ocheltree et al., 2014; compare with Mott and Peak, 2013). Although some of these models provide a pathway for incorporating the effect of ABA (Buckley, 2005), a lack of knowledge of ABA dynamics or action makes it difficult to integrate the influence of this active regulator of guard cell aperture into models. The stomatal behavior of single gene mutants (most notably the ABA synthesis and signaling mutants of Arabidopsis) strongly supports a role for ABA in mediating standard stomatal responses to changes in VPD. The stomata of these mutants are known to have less pronounced responses to a reduction in relative humidity compared with wild-type plants (Xie et al., 2006). Recently, molecular work has shown that guard cells express many of the genes required to synthesize ABA (Okamoto et al., 2009; Bauer et al., 2013), with molecular proxies for ABA level also indicating that the biochemical activity of ABA in the guard cell may increase following short-term exposure of leaves to a reduction in relative humidity (Waadt et al., 2014). These findings suggest a role for ABA in regulating stomatal responses to VPD and have led some to the conclusion that ABA synthesized autonomously by the guard cells is the predominant mechanism for stomatal responses to increased VPD (Bauer et al., 2013).Although the experimental evidence from molecular studies presents an argument for the role of ABA in the responses of stomata to changes in VPD, very few studies have quantified changes in ABA level in response to VPD. It is well established that ABA levels in leaves and guard cells can increase following the imposition of turgor loss or water stress (Pierce and Raschke, 1980; Harris et al., 1988; Harris and Outlaw, 1991). However, only a few studies have reported increases in foliar ABA level in response to high VPD (Bauerle et al., 2004; Giday et al., 2013), and none have investigated whether these observed dynamic changes or differences in ABA level were functionally relevant for stomatal control. In addition, no study has quantified the levels of ABA in guard cells during a transition in VPD.Here, we investigate the relative importance of ABA for the stomatal response to VPD in whole plants, sampled from across the vascular land plant lineage. We provide, to our knowledge, the first functional assessment of changes in ABA levels driving stomatal responses to VPD as well as critically investigate the recent suggestion that stomatal responses to VPD are driven by an autonomous guard cell synthesis of ABA.     

Response of Phytoplankton Photophysiology to Varying Environmental Conditions in the Sub-Antarctic and Polar Frontal Zone

Climate-driven changes are expected to alter the hydrography of the Sub-Antarctic Zone (SAZ) and Polar Frontal Zone (PFZ) south of Australia, in which distinct regional environments are believed to be responsible for the differences in phytoplankton biomass in these regions. Here, we report how the dynamic influences of light, iron and temperature, which are responsible for the photophysiological differences between phytoplankton in the SAZ and PFZ, contribute to the biomass differences in these regions. High effective photochemical efficiency of photosystem II (/ 0.4), maximum photosynthesis rate (), light-saturation intensity (), maximum rate of photosynthetic electron transport (1/), and low photoprotective pigment concentrations observed in the SAZ correspond to high chlorophyll and iron concentrations. In contrast, phytoplankton in the PFZ exhibits low / ( 0.2) and high concentrations of photoprotective pigments under low light environment. Strong negative relationships between iron, temperature, and photoprotective pigments demonstrate that cells were producing more photoprotective pigments under low temperature and iron conditions, and are responsible for the low biomass and low productivity measured in the PFZ. As warming and enhanced iron input is expected in this region, this could probably increase phytoplankton photosynthesis in this region. However, complex interactions between the biogeochemical processes (e.g. stratification caused by warming could prevent mixing of nutrients), which control phytoplankton biomass and productivity, remain uncertain.     

城市绿地乡土树种对大气污染的适应性响应

为探究华南乡土树种对东莞市不同绿地类型的适应性,将14种华南乡土树种分别栽植于四种绿地类型：工业绿地、道路绿地、城市公园和植物园,研究乡土树种的生理生态变化。各类型绿地的大气监测结果表明,道路绿地的各项大气污染指标明显高于其他绿地;工业区绿地的总悬浮颗粒物和氮氧化物浓度比植物园高;城市公园的总悬浮颗粒物浓度明显高于植物园。14个树种的生理生态研究结果表明,不同植物在各种绿地环境中适应策略各异,且随着污染强度的增加,多数植物净光合速率（Pn）和气孔导度（Gs）之间的相关系数降低,且Pn与Gs之间的相关系数变化不一致。14种华南乡土树种中,深山含笑、樟树、山杜英、黄栀子、胡氏青冈、九节、格木、黄果厚壳桂和华润楠等9个树种在不同污染环境下均表现出不同程度的敏感性,红鳞蒲桃和越南山龙眼对各种环境污染都不敏感,红叶石楠、假鹰爪和枫香在不同的污染环境中表现各异。     

Tree Height-Related Hydraulic Strategy to Cope with Freeze-Thaw Stress in Six Common Urban Tree Species in North China

Urban trees are sensitive to extreme weather events under climate change. Freeze-thaw induced hydraulic failure could induce urban tree dieback and nullify the services they provide. Plant height is a simple but significant trait for plant ecological strategies. Understanding how urban trees with different heights adapt to freeze-thaw stress is increasingly important under climate change. We investigated the relationship between tree height and stem hydraulic functional traits of six common urban tree species in North China to explore tree height-related hydraulic strategies to cope with freeze-thaw stress. Results showed that tall trees had wider vessels, higher hydraulic conductivity, more winter embolism, but lower vessel and wood densities. Positive relationships were found between tree height and vessel diameter, hydraulic conductivity, and freeze-thaw induced embolism, and negative relationships were found between tree height and vessel and wood densities, which implied that short trees employ more conservative ecological strategies than tall trees. Tall and short tree species were well separated by multiple stem hydraulic functional traits; this is consistent with the fact that tall and short trees occupy different niches and indicates that different hydraulic strategies for freeze-thaw stress exist between them. Tall trees might face more pressure to survive under extreme cold weather caused by climate change in the future. Therefore, more attention should be paid to tall urban tree management in North China to cope with extreme cold weather.     

Plasticity of Leaf Hydraulic Conductance in Maize in Response to Varying Nitrogen and Water Supplies

Russian Journal of Plant Physiology - The availabilities of nitrogen (N) and water are two important environmental factors affecting crop growth and productivity in the dryland farming, how they...     

Distribution of Epilithic Diatoms in Response to Environmental Conditions in an Urban Tropical Stream, Central Kenya

Use of diatoms in monitoring water quality is well acknowledged in developed countries, but only recently has the assessment started gaining importance in developing countries. Diatoms can be obtained from natural and artificial substrates. Appreciating the differences and similarities of diatom assemblages on both substrates may contribute to a better understanding and standardization particularly during monitoring of water quality. During this study we assessed diatom assemblages, biodiversity and trophic indices in relation to water quality along the Nairobi River. Fifteen sites were sampled in September 2000 during the dry season. Diatoms were collected from natural substrates (stones, pebbles) and artificial substrates (100% acrylic wool). On artificial and natural substrates, a total of 190 and 151 taxa were found, respectively, the majority of these taxa (80%) have cosmopolitan distribution and are also widespread throughout tropical African. Species composition changed downstream, five taxa dominated upper and mid stream sites whereas lower stream sites were dominated by one or two taxa. Species richness, diversity, dominance and evenness were positively correlated with NO₃, O₂ and altitude but decreased markedly downstream with a simultaneous increase in total dissolved solids, alkalinity, chemical oxygen demand and PO₄. Ordination and classification (CANOCO and TWINSPAN) showed that diatom assemblages in the Nairobi River responded strongly to water quality changes with respect to concentrations of NO₃, NO₂, total dissolved solids and temperature. Taxa common at less impacted upstream sites included Gomphonema gracilis, Anomoeoneis brachysira and Fragilaria biceps; while common taxa at midstream sites with agricultural catchments were Gomphonema parvulum, Navicula cryptocephala, N. schroeteri, N. bryophila, N. halophila, Nitzschia linearis var. linearis and Cymbella silesica. Achnanthes minutissima var. saprophila, Gomphonema angustum, Navicula subminuscula, N. arvensis, Nitzschia palea and N. umbonata were most common at urban sites, which were polluted by residential and industrial effluents. Trophic diatom indices suggested that water quality was poor at most sites in the Nairobi River. Most sites along the river had low Generic Diatom Index values, GDI (<12) and high Trophic Diatom Index values, TDI 73–78 (median = 76) and 75–84 (median = 77) for artificial and natural substrates, respectively. This study showed that diatoms' response on natural and artificial substrates were similar and reflected environmental conditions correctly.     

Analysis of Growth and Water Relations of Tomato Fruits in Relation to Air Vapor Pressure Deficit and Plant Fruit Load

The influence of air vapor pressure deficit (VPD) and plant fruit load on the expansion and water relations of young tomato fruits grown in a glasshouse were evaluated under summer Mediterranean conditions. The contributions of phloem, xylem and transpiration fluxes to the fruit volume increase were estimated at an hourly scale from the growth curves of intact, heat-girdled and detached fruits, measured using displacement transducers. High VPD conditions reduced the xylem influx and increased the fruit transpiration, but hardly affected the phloem influx. Net water accumulation and growth rate were reduced, and a xylem efflux even occurred during the warmest and driest hours of the day. Changes in xylem flux could be explained by variations in the gradient of water potential between stem and fruit, due to changes in stem water potential. Misting reduced air VPD and alleviated the reduction in fruit volume increase through an increase in xylem influx and a decrease in fruit transpiration. Under low fruit load, the competition for assimilates being likely reduced, the phloem flux to fruits increased, similarly to the xylem and transpiration fluxes, without any changes in the fruit water potential. However, different diurnal dynamics among treatments assume variable contributions of turgor and osmotic pressure in F3 and F6 fruits, and hypothetical short-term variations in the water potential gradient between stem and fruit, preventing xylem efflux in F3 fruits.     

云冷杉天然林乔木树种组成及物种多样性对择伐强度的动态响应

以吉林省长白山金沟岭林场云冷杉天然林为研究对象,定量分析不同择伐强度(对照0%、弱度择伐20.6%、中度择伐29.9%和重度择伐41.6%)下其乔木层树种组成及物种多样性的动态变化规律。结果显示:采伐后经10年恢复,择伐对原有林分结构影响不大,其中弱度和中度择伐仍能保持择伐前以冷杉占优势、红松和云杉占亚优势的物种结构,重度择伐使林分结构略有改变,但不明显;相比择伐前,择伐后经10年恢复的样地α多样性指数均有所增加,优势度指数均有不同程度的降低,其中弱度择伐样地的物种丰富度指数、多样性指数及均匀度指数的增加量和优势度指数的减少量均最大,说明弱度择伐有利于改善林分物种多样性状况,使林分物种更为丰富。本研究结果表明采取弱度择伐的方式更有利于维持林分物种结构及物种多样性,有利于云冷杉天然林的可持续经营和发展。     

Response Breadths on Environmental Gradients of Germination and Seedling Growth in Two Dominant Forest Tree Species of Central Himalaya

The responses of Pimis roxburghii (chir pine), an early successionalspecies, andQuercus leucotrichophora(banj oak), a late successionalspecies were compared in terms of seed germination and seedlinggrowth. Seed germination was observed on single-factor gradientsof temperature, water stress, seed desiccation and light, andseedling growth on gradients of shade and soil moisture. Thesuppression of germination caused by continuous dark and far-redwas greater in Pinus than in Quercus. Pinus seed germinationshowed wider response breadth on gradients of water stress andseed desiccation, and narrower response breadth on the gradientof light quality, compared withQuercus. In terms of seedlinggrowth, the response breadth of Pinus was wider on the gradientof soil moisture and narrower on the gradient of shade comparedwith Quercus. Under higher soil moisture stress, the seedlingsof both species attained similar heights, butQuercus seedlingsachieved higher dry weights, a higher root: shoot ratio andlower leaf weight ratio than Pinus.     

Hydraulic Conductance and Stomatal Sensitivity to Changes of Leaf Water Status in Six Deciduous Tree Species

The relationship between shoot hydraulic conductance (L) and stomatal sensitivity to changes in leaf water status was studied in the saplings of six deciduous tree species. L increased significantly in sequence: Acer platanoides < Tilia Cordata < Padus avium = Quercus robur < Salix caprea = Populus tremula. L was higher in the trees grown in soil with a higher nitrogen content and lower in the trees grown under mild water stress or kept in darkness for several days. L was higher in July than in September in all the species. L correlated positively with maximum photosynthesis, stomatal conductance and stomatal sensitivity to an increase in leaf water potential, but negatively with stomatal sensitivity to a decrease in leaf water potential. The correlations between L and any other parameter were approximated by three different curves: data for water-stressed plants fit to the first, data for plants kept in darkness fit to the second and all the other data fit to the third curve. The reasons of the differences of shoot hydraulic conductance in the different experimental sets and the mechanisms which may cause the correlation between L and the other characteristics are discussed.     

Adaptations of the Secretome of Candida albicans in Response to Host-Related Environmental Conditions

The wall proteome and the secretome of the fungal pathogen Candida albicans help it to thrive in multiple niches of the human body. Mass spectrometry has allowed researchers to study the dynamics of both subproteomes. Here, we discuss some major responses of the secretome to host-related environmental conditions. Three β-1,3-glucan-modifying enzymes, Mp65, Sun41, and Tos1, are consistently found in large amounts in culture supernatants, suggesting that they are needed for construction and expansion of the cell wall β-1,3-glucan layer and thus correlate with growth and might serve as diagnostic biomarkers. The genes ENG1, CHT3, and SCW11, which encode an endoglucanase, the major chitinase, and a β-1,3-glucan-modifying enzyme, respectively, are periodically expressed and peak in M/G₁. The corresponding protein abundances in the medium correlate with the degree of cell separation during single-yeast-cell, pseudohyphal, and hyphal growth. We also discuss the observation that cells treated with fluconazole, or other agents causing cell surface stress, form pseudohyphal aggregates. Fluconazole-treated cells secrete abundant amounts of the transglucosylase Phr1, which is involved in the accumulation of β-1,3-glucan in biofilms, raising the question whether this is a general response to cell surface stress. Other abundant secretome proteins also contribute to biofilm formation, emphasizing the important role of secretome proteins in this mode of growth. Finally, we discuss the relevance of these observations to therapeutic intervention. Together, these data illustrate that C. albicans actively adapts its secretome to environmental conditions, thus promoting its survival in widely divergent niches of the human body.     

Metabolic and Regulatory Responses in Citrus Rootstocks in Response to Adverse Environmental Conditions

In response to adverse environmental conditions, plants modify their metabolism to adapt to the new conditions. To differentiate common responses to abiotic stress from specific adaptation to a certain stress condition, two citrus rootstocks (Carrizo citrange and Cleopatra mandarin) with a different ability to tolerate stress were subjected to soil flooding and drought, two water stress conditions. In response to these conditions, both genotypes showed altered root proline and phenylpropanoid levels, especially cinnamic acid, which was a common feature to Carrizo and Cleopatra. This was correlated with alterations in the levels of phenylpropanoid derivatives likely involved in lignin biosynthesis. In the regulatory part, levels of both stress hormones abscisic acid (ABA) and jasmonic acid (JA) decreased in response to soil flooding irrespective of the genotype’s relative flooding tolerance, but, on the other hand, the concentration of both metabolites increased in response to drought, showing a transient accumulation of JA after a few days and a progressive pattern of ABA increase. These responses are probably associated with different regulatory processes under soil flooding and drought. In addition, alterations in indole acetic acid (IAA) levels in citrus roots seemed to be associated with particular stress tolerance. Moreover, both genotypes exhibited a low degree of overlap in the metabolites induced under similar stress conditions, indicating a specific mechanism to cope with stress in plant species. Results also indicated a different metabolic basal status in both genotypes that could contribute to stress tolerance.     

四种热带雨林树种光合和形态解剖特征对不同生长光强的适应

研究了西双版纳热带雨林2种喜光树种中平树(Macaranga denticulata)、倒樱木(Pnravallaris macrophylla)和2种耐荫树种云南肉豆蔻(Myristica yunnanensis)、金丝李(Garcinia paucinervis)幼苗叶片光合和形态解剖特征对3种不同生长光强(5％、25%和50％相对光强)的适应。研究结果表明,与强光下相比,弱光下生长的4种植物最大净光合速率、光饱和点、光补偿点、暗呼吸速率、叶绿素a／b、叶片和栅栏组织厚度、气孔密度和比叶重都降低,而海绵组织／栅栏组织和叶绿素含量升高。在相同光强下,与2种耐荫树种相比,2种喜光树种有较大的最大净光合速率、暗呼吸速率、气孔密度和较低的叶绿素含量。在不同光强下,4种植物均表现出了对光适应有利的生理和形态解剖可塑性,而喜光树种比耐荫树种有较大的生理和形态可塑性,表明喜光树种具有比耐荫树种对强光有更强的适应能力。4种植物的生理指标的可塑性均大于叶片解剖结构的可塑性。     

Control of Photosynthesis and Stomatal Conductance in Ricinus communis L. (Castor Bean) by Leaf to Air Vapor Pressure Deficit

Castor bean (Ricinus communis L.) has a high photosynthetic capacity under high humidity and a pronounced sensitivity of photosynthesis to high water vapor pressure deficit (VPD). The sensitivity of photosynthesis to varying VPD was analyzed by measuring CO₂ assimilation, stomatal conductance (g_s), quantum yield of photosystem II (_II), and nonphotochemical quenching of chlorophyll fluorescence (q_N) under different VPD. Under both medium (1000) and high (1800 micromoles quanta per square meter per second) light intensities, CO₂ assimilation decreased as the VPD between the leaf and the air around the leaf increased. The g_s initially dropped rapidly with increasing VPD and then showed a slower decrease above a VPD of 10 to 20 millibars. Over a temperature range from 20 to 40°C, CO₂ assimilation and g_s were inhibited by high VPD (20 millibars). However, the rate of transpiration increased with increasing temperature at either low or high VPD due to an increase in g_s. The relative inhibition of photosynthesis under photorespiring (atmospheric levels of CO₂ and O₂) versus nonphotorespiring (700 microbars CO₂ and 2% O₂) conditions was greater under high VPD (30 millibars) than under low VPD (3 millibars). Also, with increasing light intensity the relative inhibition of photosynthesis by O₂ increased under high VPD, but decreased under low VPD. The effect of high VPD on photosynthesis under various conditions could not be totally accounted for by the decrease in the intercellular CO₂ in the leaf (C_i) where C_i was estimated from gas exchange measurements. However, estimates of C_i from measurements of _II and q_N suggest that the decrease in photosynthesis and increase in photorespiration under high VPD can be totally accounted for by stomatal closure and a decrease in C_i. The results also suggest that nonuniform closure of stomata may occur in well-watered plants under high VPD, causing overestimates in the calculation of C_i from gas exchange measurements. Under low VPD, 30°C, high light, and saturating CO₂, castor bean (C₃ tropical shrub) has a rate of photosynthesis (61 micromoles CO₂ per square meter per second) that is about 50% higher than that of tobacco (C₃) or maize (C₄) under the same conditions. The chlorophyll content, total soluble protein, and ribulose-1,5-bisphosphate carboxylase/oxygenase level on a leaf area basis were much higher in castor bean than in maize or tobacco, which accounts for its high rates of photosynthesis under low VPD.     

Controlled Environment Chambers for Investigating Tree Response to Elevated CO2 and Temperature Under Boreal Conditions

A closed CO₂ and temperature-controlled, long-term chamber system has been developed and set up in a typical boreal forest of Scots pine (Pinus sylvestris L.) near the Mekrijärvi Research Station (62°47N, 30°58E, 145 m above sea level) belonging to the University of Joensuu, Finland. The main objectives of the experiment were to provide a means of assessing the medium to long-term effects of elevated atmospheric CO₂ concentration (EC) and temperature (ET) on photosynthesis, respiration, growth, and biomass at the whole-tree level and to measure instantaneous whole-system CO₂ exchange. The system consists of 16 chambers with individual facilities for controlling CO₂ concentration, temperature, and the combination of the two. The chambers can provide a wide variety of climatic conditions that are similar to natural regimes. In this experiment the target CO₂ concentration in the EC chambers was set at a fixed constant of 700 µmol mol^–1 and the target air temperature in the ET chambers to track the ambient temperature but with a specified addition. Chamber performance was assessed on the base of recordings covering three consecutive years. The CO₂ and temperature control in these closed chambers was in general accurate and reliable. CO₂ concentration in the EC chambers was within 600–725 µmol mol^–1 for 90 % of the exposure time during the "growing-season" (15 April – 15 September) and 625–725 µmol mol^–1 for 88 % of the time in the "off-season" (16 September – 14 April), while temperatures in the chambers were within ±2.0 °C of the ambient or target temperature in the "growing season" and within ±3.0 °C in the "off season". There were still some significant chamber effects. Solar radiation in the chambers was reduced by 50–60 % for 82 % of the time in the "growing season" and 55–65 % for 78 % of the time in the "off season", and the relative humidity of the air was increased by 5–10 % for 72 % of the time in the "growing season" and 2–12 % for 91 % of the time in the "off season". The crown architecture and main phenophase of the trees were not modified significantly by enclosure in the chambers, but some physiological parameters changed significantly, e.g., the radiant energy-saturated photosynthesis rate, transpiration rate, maximum photochemical efficiency of photosystem 2, and chlorophyll content.     

Species-Independent Down-Regulation of Leaf Photosynthesis and Respiration in Response to Shading: Evidence from Six Temperate Tree Species

The ability to down-regulate leaf maximum net photosynthetic capacity (Amax) and dark respiration rate (Rdark) in response to shading is thought to be an important adaptation of trees to the wide range of light environments that they are exposed to across space and time. A simple, general rule that accurately described this down-regulation would improve carbon cycle models and enhance our understanding of how forest successional diversity is maintained. In this paper, we investigated the light response of Amax and Rdark for saplings of six temperate forest tree species in New Jersey, USA, and formulated a simple model of down-regulation that could be incorporated into carbon cycle models. We found that full-sun values of Amax and Rdark differed significantly among species, but the rate of down-regulation (proportional decrease in Amax or Rdark relative to the full-sun value) in response to shade was not significantly species- or taxon-specific. Shade leaves of sun-grown plants appear to follow the same pattern of down-regulation in response to shade as leaves of shade-grown plants. Given the light level above a leaf and one species-specific number (either the full-sun Amax or full-sun Rdark), we provide a formula that can accurately predict the leaf''s Amax and Rdark. We further show that most of the down regulation of per unit area Rdark and Amax is caused by reductions in leaf mass per unit area (LMA): as light decreases, leaves get thinner, while per unit mass Amax and Rdark remain approximately constant.     

Photosynthesis Decrease and Stomatal Control of Gas Exchange in Abies alba Mill. in Response to Vapor Pressure Difference

The responses of steady state CO₂ assimilation rate (A), transpiration rate (E), and stomatal conductance (g_s) to changes in leaf-to-air vapor pressure difference (ΔW) were examined on different dates in shoots from Abies alba trees growing outside. In Ecouves, a provenance representative of wet oceanic conditions in Northern France, both A and g_s decreased when ΔW was increased from 4.6 to 14.5 Pa KPa⁻¹. In Nebias, which represented the dry end of the natural range of A. alba in southern France, A and g_s decreased only after reaching peak levels at 9.0 and 7.0 Pa KPa⁻¹, respectively. The representation of the data in assimilation rate (A) versus intercellular CO₂ partial pressure (C_i) graphs allowed us to determine how stomata and mesophyll photosynthesis interacted when ΔW was increased. Changes in A were primarily due to alterations in mesophyll photosynthesis. At high ΔW, and especially in Ecouves when soil water deficit prevailed, A declined, while C_i remained approximately constant, which may be interpreted as an adjustment of g_s to changes in mesophyll photosynthesis. Such a stomatal control of gas exchange appeared as an alternative to the classical feedforward interpretation of E versus ΔW responses with a peak rate of E. The gas exchange response to ΔW was also characterized by considerable deviations from the optimization theory of IR Cowan and GD Farquhar (1977 Symp Soc Exp Biol 31: 471-505).