A model of canopy photosynthesis incorporating protein distribution through the canopy and its acclimation to light, temperature and CO2

Background and Aims

The distribution of photosynthetic enzymes, or nitrogen, through the canopy affects canopy photosynthesis, as well as plant quality and nitrogen demand. Most canopy photosynthesis models assume an exponential distribution of nitrogen, or protein, through the canopy, although this is rarely consistent with experimental observation. Previous optimization schemes to derive the nitrogen distribution through the canopy generally focus on the distribution of a fixed amount of total nitrogen, which fails to account for the variation in both the actual quantity of nitrogen in response to environmental conditions and the interaction of photosynthesis and respiration at similar levels of complexity.

Model

A model of canopy photosynthesis is presented for C₃ and C₄ canopies that considers a balanced approach between photosynthesis and respiration as well as plant carbon partitioning. Protein distribution is related to irradiance in the canopy by a flexible equation for which the exponential distribution is a special case. The model is designed to be simple to parameterize for crop, pasture and ecosystem studies. The amount and distribution of protein that maximizes canopy net photosynthesis is calculated.

Key Results

The optimum protein distribution is not exponential, but is quite linear near the top of the canopy, which is consistent with experimental observations. The overall concentration within the canopy is dependent on environmental conditions, including the distribution of direct and diffuse components of irradiance.

Conclusions

The widely used exponential distribution of nitrogen or protein through the canopy is generally inappropriate. The model derives the optimum distribution with characteristics that are consistent with observation, so overcoming limitations of using the exponential distribution. Although canopies may not always operate at an optimum, optimization analysis provides valuable insight into plant acclimation to environmental conditions. Protein distribution has implications for the prediction of carbon assimilation, plant quality and nitrogen demand.     

The impact of modifying photosystem antenna size on canopy photosynthetic efficiency—Development of a new canopy photosynthesis model scaling from metabolism to canopy level processes

Canopy photosynthesis (A_c) describes photosynthesis of an entire crop field and the daily and seasonal integrals of A_c positively correlate with daily and seasonal biomass production. Much effort in crop breeding has focused on improving canopy architecture and hence light distribution inside the canopy. Here, we develop a new integrated canopy photosynthesis model including canopy architecture, a ray tracing algorithm, and C₃ photosynthetic metabolism to explore the option of manipulating leaf chlorophyll concentration ([Chl]) for greater A_c and nitrogen use efficiency (NUE). Model simulation results show that (a) efficiency of photosystem II increased when [Chl] was decreased by decreasing antenna size and (b) the light received by leaves at the bottom layers increased when [Chl] throughout the canopy was decreased. Furthermore, the modelling revealed a modest ~3% increase in A_c and an ~14% in NUE was accompanied when [Chl] reduced by 60%. However, if the leaf nitrogen conserved by this decrease in leaf [Chl] were to be optimally allocated to other components of photosynthesis, both A_c and NUE can be increased by over 30%. Optimizing [Chl] coupled with strategic reinvestment of conserved nitrogen is shown to have the potential to support substantial increases in A_c, biomass production, and crop yields.     

Increasing canopy photosynthesis in rice can be achieved without a large increase in water use—A model based on free‐air CO2 enrichment

Achieving higher canopy photosynthesis rates is one of the keys to increasing future crop production; however, this typically requires additional water inputs because of increased water loss through the stomata. Lowland rice canopies presently consume a large amount of water, and any further increase in water usage may significantly impact local water resources. This situation is further complicated by changing the environmental conditions such as rising atmospheric CO₂ concentration ([CO₂]). Here, we modeled and compared evapotranspiration of fully developed rice canopies of a high‐yielding rice cultivar (Oryza sativa L. cv. Takanari) with a common cultivar (cv. Koshihikari) under ambient and elevated [CO₂] (A‐CO₂ and E‐CO₂, respectively) via leaf ecophysiological parameters derived from a free‐air CO₂ enrichment (FACE) experiment. Takanari had 4%–5% higher evapotranspiration than Koshihikari under both A‐CO₂ and E‐CO₂, and E‐CO₂ decreased evapotranspiration of both varieties by 4%–6%. Therefore, if Takanari was cultivated under future [CO₂] conditions, the cost for water could be maintained at the same level as for cultivating Koshihikari at current [CO₂] with an increase in canopy photosynthesis by 36%. Sensitivity analyses determined that stomatal conductance was a significant physiological factor responsible for the greater canopy photosynthesis in Takanari over Koshihikari. Takanari had 30%–40% higher stomatal conductance than Koshihikari; however, the presence of high aerodynamic resistance in the natural field and lower canopy temperature of Takanari than Koshihikari resulted in the small difference in evapotranspiration. Despite the small difference in evapotranspiration between varieties, the model simulations showed that Takanari clearly decreased canopy and air temperatures within the planetary boundary layer compared to Koshihikari. Our results indicate that lowland rice varieties characterized by high‐stomatal conductance can play a key role in enhancing productivity and moderating heat‐induced damage to grain quality in the coming decades, without significantly increasing crop water use.     

Plant growth and respiration re-visited: maintenance respiration defined - it is an emergent property of, not a separate process within, the system - and why the respiration : photosynthesis ratio is conservative

Background and Aims

Plant growth and respiration still has unresolved issues, examined here using a model. The aims of this work are to compare the model''s predictions with McCree''s observation-based respiration equation which led to the ‘growth respiration/maintenance respiration paradigm’ (GMRP) – this is required to give the model credibility; to clarify the nature of maintenance respiration (MR) using a model which does not represent MR explicitly; and to examine algebraic and numerical predictions for the respiration:photosynthesis ratio.

Methods

A two-state variable growth model is constructed, with structure and substrate, applicable on plant to ecosystem scales. Four processes are represented: photosynthesis, growth with growth respiration (GR), senescence giving a flux towards litter, and a recycling of some of this flux. There are four significant parameters: growth efficiency, rate constants for substrate utilization and structure senescence, and fraction of structure returned to the substrate pool.

Key Results

The model can simulate McCree''s data on respiration, providing an alternative interpretation to the GMRP. The model''s parameters are related to parameters used in this paradigm. MR is defined and calculated in terms of the model''s parameters in two ways: first during exponential growth at zero growth rate; and secondly at equilibrium. The approaches concur. The equilibrium respiration:photosynthesis ratio has the value of 0·4, depending only on growth efficiency and recycling fraction.

Conclusions

McCree''s equation is an approximation that the model can describe; it is mistaken to interpret his second coefficient as a maintenance requirement. An MR rate is defined and extracted algebraically from the model. MR as a specific process is not required and may be replaced with an approach from which an MR rate emerges. The model suggests that the respiration:photosynthesis ratio is conservative because it depends on two parameters only whose values are likely to be similar across ecosystems.     

黄瓜单叶净光合速率对二氧化碳浓度、温度和光照强度响应模型

以日光温室黄瓜品种津优一号和露地品种津研四号为试材，在人工气候室内测定了特定环境条件（CO₂浓度、光照强度和温度等）下的黄瓜单叶净光合速率（P_n），结合数学方法，建立了模拟黄瓜单叶P_n(y)对CO₂浓度(x₁)、光照强度(x₂)和温度(x₃)3因子响应的数学模型: 津优一号y=exp(-242.1217/x₁)［61.0202-0.11(x₂-30.926)²］exp(-272.8874/x₃)+0.9355；津研四号y=exp(-179.8803/x₁)［50.0771-0.0609(x₂-34.3455)²］exp(-267.9653/x₃)+0.7377.由模型可知，P_n对温度的响应为二次曲线，对CO₂浓度和光照强度的响应为指数函数.     

Diurnal changes in leaf gas exchange and validation of a mathematical model for coffee (<Emphasis Type="Italic">Coffea arabica</Emphasis> L.) canopy photosynthesis

Diurnal variations in net photosynthetic rate (P _N), transpiration rate (E), stomatal conductance (g _s), internal CO₂ concentration (C _i), and water use efficiency (WUE) were studied on individual leaves of coffee plants to determine the effect of climatic factors on photosynthetic capacity. P _N and E showed bimodal behaviour with the maximum values of P _N at mid-morning. At noon, under saturating photosynthetically active radiation (PAR) and high leaf temperature (T _l), P _N declined. In the afternoon (14:00), P _N slightly recovered in association with a decrease in Tl and in leaf-to-air vapour pressure deficit (VPD). Reductions in E during the morning were associated with decreases in g _s. Higher WUE in the morning was related to higher P _N and lower E. The reverse occurred in the afternoon. Goudriaan's simulation model, adapted for coffee canopy photosynthesis, was tested at the level of whole plant (P _pl). Three methods were used: (a) Whole plant net photosynthesis (P _pl) under semi-controlled conditions in a chamber. (b) P _pl estimation following Goudriaan's method (Gaussian integration) of instantaneous P _N in single leaves at three canopy depths and at three different hours assuming a photosynthesis unimodal behaviour. (c) P _pl using Goudriaan's method but at five different hours according to the bimodal behaviour reported above. Results of P _pl estimates using Goudriaan's model adapted for coffee canopy confirm the observed P _pl bimodal behaviour with high fitness degree of the measured whole plant photosynthesis. The high fitness found among observed and simulated data indicates that the modified model may be used as a subroutine for the general simulation model of coffee crop growth.     

Climatic controls on the carbon and water balances of a boreal aspen forest, 1994–2003

The carbon and water budgets of boreal and temperate broadleaf forests are sensitive to interannual climatic variability and are likely to respond to climate change. This study analyses 9 years of eddy‐covariance data from the Boreal Ecosystem Research and Monitoring Sites (BERMS) Southern Old Aspen site in central Saskatchewan, Canada and characterizes the primary climatic controls on evapotranspiration, net ecosystem production (F_NEP), gross ecosystem photosynthesis (P) and ecosystem respiration (R). The study period was dominated by two climatic extremes: extreme warm and cool springs, which produced marked contrasts in the canopy duration, and a severe, 3‐year drought. Annual F_NEP varied among years from 55 to 367 g C m⁻² (mean 172, SD 94). Interannual variability in F_NEP was controlled primarily by factors that affected the R/P ratio, which varied between 0.74 and 0.96 (mean 0.87, SD 0.06). Canopy duration enhanced P and F_NEP with no apparent effect on R. The fraction of annual photosynthetically active radiation (PAR) that was absorbed by the canopy foliage varied from 38% in late leaf‐emergence years to 51% in early leaf‐emergence years. Photosynthetic light‐use efficiency (mean 0.0275, SD 0.026 mol C mol⁻¹ photons) was relatively constant during nondrought years but declined with drought intensity to a minimum of 0.0228 mol C mol⁻¹ photons during the most severe drought year. The impact of drought on F_NEP varied with drought intensity. Years of mild‐to‐moderate drought suppressed R while having little effect on P, so that F_NEP was enhanced. Years of severe drought suppressed both R and P, causing either little change or a subtle reduction in F_NEP. The analysis produced new insights into the dominance of canopy duration as the most important biophysical control on F_NEP. The results suggested a simple conceptual model for annual F_NEP in boreal deciduous forests. When water is not limiting, annual P is controlled by canopy duration via its influence on absorbed PAR at constant light‐use efficiency. Water stress suppresses P, by reducing light‐use efficiency, and R, by limiting growth and/or suppressing microbial respiration. The high photosynthetic light‐use efficiency showed this site to be a highly productive boreal deciduous forest, with properties similar to many temperate deciduous forests.     

Lower responsiveness of canopy evapotranspiration rate than of leaf stomatal conductance to open‐air CO2 elevation in rice

An elevated atmospheric CO₂ concentration ([CO₂]) can reduce stomatal conductance of leaves for most plant species, including rice (Oryza sativa L.). However, few studies have quantified seasonal changes in the effects of elevated [CO₂] on canopy evapotranspiration, which integrates the response of stomatal conductance of individual leaves with other responses, such as leaf area expansion, changes in leaf surface temperature, and changes in developmental stages, in field conditions. We conducted a field experiment to measure seasonal changes in stomatal conductance of the uppermost leaves and in the evapotranspiration, transpiration, and evaporation rates using a lysimeter method. The study was conducted for flooded rice under open‐air CO₂ elevation. Stomatal conductance decreased by 27% under elevated [CO₂], averaged throughout the growing season, and evapotranspiration decreased by an average of 5% during the same period. The decrease in daily evapotranspiration caused by elevated [CO₂] was more significantly correlated with air temperature and leaf area index (LAI) rather than with other parameters of solar radiation, days after transplanting, vapor‐pressure deficit and FAO reference evapotranspiration. This indicates that higher air temperatures, within the range from 16 to 27 °C, and a larger LAI, within the range from 0 to 4 m² m^?2, can increase the magnitude of the decrease in evapotranspiration rate caused by elevated [CO₂]. The crop coefficient (i.e. the evapotranspiration rate divided by the FAO reference evapotranspiration rate) was 1.24 at ambient [CO₂] and 1.17 at elevated [CO₂]. This study provides the first direct measurement of the effects of elevated [CO₂] on rice canopy evapotranspiration under open‐air conditions using the lysimeter method, and the results will improve future predictions of water use in rice fields.     

Responses of terrestrial ecosystems to temperature and precipitation change: a meta‐analysis of experimental manipulation

Global mean temperature is predicted to increase by 2–7 °C and precipitation to change across the globe by the end of this century. To quantify climate effects on ecosystem processes, a number of climate change experiments have been established around the world in various ecosystems. Despite these efforts, general responses of terrestrial ecosystems to changes in temperature and precipitation, and especially to their combined effects, remain unclear. We used meta‐analysis to synthesize ecosystem‐level responses to warming, altered precipitation, and their combination. We focused on plant growth and ecosystem carbon (C) balance, including biomass, net primary production (NPP), respiration, net ecosystem exchange (NEE), and ecosystem photosynthesis, synthesizing results from 85 studies. We found that experimental warming and increased precipitation generally stimulated plant growth and ecosystem C fluxes, whereas decreased precipitation had the opposite effects. For example, warming significantly stimulated total NPP, increased ecosystem photosynthesis, and ecosystem respiration. Experimentally reduced precipitation suppressed aboveground NPP (ANPP) and NEE, whereas supplemental precipitation enhanced ANPP and NEE. Plant productivity and ecosystem C fluxes generally showed higher sensitivities to increased precipitation than to decreased precipitation. Interactive effects of warming and altered precipitation tended to be smaller than expected from additive, single‐factor effects, though low statistical power limits the strength of these conclusions. New experiments with combined temperature and precipitation manipulations are needed to conclusively determine the importance of temperature–precipitation interactions on the C balance of terrestrial ecosystems under future climate conditions.     

Behavioral signature of intraspecific competition and density dependence in colony‐breeding marine predators

In populations of colony‐breeding marine animals, foraging around colonies can lead to intraspecific competition. This competition affects individual foraging behavior and can cause density‐dependent population growth. Where behavioral data are available, it may be possible to infer the mechanism of intraspecific competition. If these mechanics are understood, they can be used to predict the population‐level functional response resulting from the competition. Using satellite relocation and dive data, we studied the use of space and foraging behavior of juvenile and adult gray seals (Halichoerus grypus) from a large (over 200,000) and growing population breeding at Sable Island, Nova Scotia (44.0 ^oN 60.0 ^oW). These data were first analyzed using a behaviorally switching state‐space model to infer foraging areas followed by randomization analysis of foraging region overlap of competing age classes. Patterns of habitat use and behavioral time budgets indicate that young‐of‐year juveniles (YOY) were likely displaced from foraging areas near (<10 km) the breeding colony by adult females. This displacement was most pronounced in the summer. Additionally, our data suggest that YOY are less capable divers than adults and this limits the habitat available to them. However, other segregating mechanisms cannot be ruled out, and we discuss several alternate hypotheses. Mark–resight data indicate juveniles born between 1998 and 2002 have much reduced survivorship compared with cohorts born in the late 1980s, while adult survivorship has remained steady. Combined with behavioral observations, our data suggest YOY are losing an intraspecific competition between adults and juveniles, resulting in the currently observed decelerating logistic population growth. Competition theory predicts that intraspecific competition resulting in a clear losing competitor should cause compensatory population regulation. This functional response produces a smooth logistic growth curve as carrying capacity is approached, and is consistent with census data collected from this population over the past 50 years. The competitive mechanism causing compensatory regulation likely stems from the capital‐breeding life‐history strategy employed by gray seals. This strategy decouples reproductive success from resources available around breeding colonies and prevents females from competing with each other while young are dependent.     

Functional diversity of photosynthesis during drought in a model tropical rainforest – the contributions of leaf area, photosynthetic electron transport and stomatal conductance to reduction in net ecosystem carbon exchange

The tropical rainforest mesocosm within the Biosphere 2 Laboratory, a model system of some 110 species developed over 12 years under controlled environmental conditions, has been subjected to a series of comparable drought experiments during 2000–2002. In each study, the mesocosm was subjected to a 4–6 week drought, with well‐defined rainfall events before and after the treatment. Ecosystem CO₂ uptake rate (A_eco) declined 32% in response to the drought, with changes occurring within days and being reversible within weeks, even though the deeper soil layers did not become significantly drier and leaf‐level water status of most large trees was not greatly affected. The reduced A_eco during the drought reflected both morphological and physiological responses. It is estimated that the drought‐induced 32% reduction of A_eco has three principal components: (1) leaf fall increased two‐fold whereas leaf expansion growth of some canopy dominants declined to 60%, leading to a 10% decrease in foliage coverage of the canopy. This might be the main reason for the persistent reduction of A_eco after rewatering. (2) The maximum photosynthetic electron transport rate at high light intensities in remaining leaves was reduced to 71% for three of the four species measured, even though no chronic photo‐inhibition occurred. (3) Stomata closed, leading to a reduced ecosystem water conductance to water vapour (33% of pre‐drought values), which not only reduced ecosystem carbon uptake rate, but may also have implications for water and energy budgets of tropical ecosystems. Additionally, individual rainforest trees responded differently, expressing different levels of stress and stress avoiding mechanisms. This functional diversity renders the individual response heterogeneous and has fundamental implications to scale leaf level responses to ecosystem dynamics.     

The relationship of leaf photosynthetic traits – Vcmax and Jmax – to leaf nitrogen,leaf phosphorus,and specific leaf area: a meta‐analysis and modeling study

Great uncertainty exists in the global exchange of carbon between the atmosphere and the terrestrial biosphere. An important source of this uncertainty lies in the dependency of photosynthesis on the maximum rate of carboxylation (V_cmax) and the maximum rate of electron transport (J_max). Understanding and making accurate prediction of C fluxes thus requires accurate characterization of these rates and their relationship with plant nutrient status over large geographic scales. Plant nutrient status is indicated by the traits: leaf nitrogen (N), leaf phosphorus (P), and specific leaf area (SLA). Correlations between V_cmax and J_max and leaf nitrogen (N) are typically derived from local to global scales, while correlations with leaf phosphorus (P) and specific leaf area (SLA) have typically been derived at a local scale. Thus, there is no global-scale relationship between V_cmax and J_max and P or SLA limiting the ability of global-scale carbon flux models do not account for P or SLA. We gathered published data from 24 studies to reveal global relationships of V_cmax and J_max with leaf N, P, and SLA. V_cmax was strongly related to leaf N, and increasing leaf P substantially increased the sensitivity of V_cmax to leaf N. J_max was strongly related to V_cmax, and neither leaf N, P, or SLA had a substantial impact on the relationship. Although more data are needed to expand the applicability of the relationship, we show leaf P is a globally important determinant of photosynthetic rates. In a model of photosynthesis, we showed that at high leaf N (3 gm⁻²), increasing leaf P from 0.05 to 0.22 gm⁻² nearly doubled assimilation rates. Finally, we show that plants may employ a conservative strategy of J_max to V_cmax coordination that restricts photoinhibition when carboxylation is limiting at the expense of maximizing photosynthetic rates when light is limiting.     

The acclimation of leaf photosynthesis of wheat and rice to seasonal temperature changes in T‐FACE environments

Crops show considerable capacity to adjust their photosynthetic characteristics to seasonal changes in temperature. However, how photosynthesis acclimates to changes in seasonal temperature under future climate conditions has not been revealed. We measured leaf photosynthesis (A_n) of wheat (Triticum aestivum L.) and rice (Oryza sativa L.) grown under four combinations of two levels of CO₂ (ambient and enriched up to 500 µmol/mol) and two levels of canopy temperature (ambient and increased by 1.5–2.0°C) in temperature by free‐air CO₂ enrichment (T‐FACE) systems. Parameters of a biochemical C₃‐photosynthesis model and of a stomatal conductance (g_s) model were estimated for the four conditions and for several crop stages. Some biochemical parameters related to electron transport and most g_s parameters showed acclimation to seasonal growth temperature in both crops. The acclimation response did not differ much between wheat and rice, nor among the four treatments of the T‐FACE systems, when the difference in the seasonal growth temperature was accounted for. The relationships between biochemical parameters and leaf nitrogen content were consistent across leaf ranks, developmental stages, and treatment conditions. The acclimation had a strong impact on g_s model parameters: when parameter values of a particular stage were used, the model failed to correctly estimate g_s values of other stages. Further analysis using the coupled g_s–biochemical photosynthesis model showed that ignoring the acclimation effect did not result in critical errors in estimating leaf photosynthesis under future climate, as long as parameter values were measured or derived from data obtained before flowering.     

Combined effects of light and nitrate supplies on the growth,photosynthesis and ultraviolet‐absorbing compounds in marine macroalga Gracilaria lemaneiformis (Rhodophyta), with special reference to the effects of solar ultraviolet radiation

It is well known that light and nutrients are essential to plants; however, there are few investigations in which these have been studied in combination on macroalgae, especially when solar ultraviolet radiation (UVR) is concerned. We cultured the red alga Gracilaria lemaneiformis (Bory) at different nitrate concentrations and light levels with or without UVR for 24 days. The results showed that nitrate supply markedly enhanced the growth and photosynthesis, increased the absorptivity of UV‐absorbing compounds (UVACs), and decreased photoinhibition in the presence of UVR. The thalli that received photosynthetically active radiation (PAR) treatment exhibited higher growth rates than those that received PAR + UVR at ambient or enhanced nitrate concentrations. However, under PAR + UVR treatment, the absorptivity of UVACs was higher than that of PAR and fluctuated with light levels. UVR was found to reduce the maximal net photosynthetic rate, apparent photosynthetic efficiency and light‐saturating irradiance while increasing the dark respiration rate, and inducing higher inhibition of growth and photosynthesis under high light versus under low light. Ultraviolet B significantly induced the synthesis of UVACs but led to higher inhibition on growth and photosynthesis than ultraviolet A.     

Range position and climate sensitivity: The structure of among‐population demographic responses to climatic variation

Species’ distributions will respond to climate change based on the relationship between local demographic processes and climate and how this relationship varies based on range position. A rarely tested demographic prediction is that populations at the extremes of a species’ climate envelope (e.g., populations in areas with the highest mean annual temperature) will be most sensitive to local shifts in climate (i.e., warming). We tested this prediction using a dynamic species distribution model linking demographic rates to variation in temperature and precipitation for wood frogs (Lithobates sylvaticus) in North America. Using long‐term monitoring data from 746 populations in 27 study areas, we determined how climatic variation affected population growth rates and how these relationships varied with respect to long‐term climate. Some models supported the predicted pattern, with negative effects of extreme summer temperatures in hotter areas and positive effects on recruitment for summer water availability in drier areas. We also found evidence of interacting temperature and precipitation influencing population size, such as extreme heat having less of a negative effect in wetter areas. Other results were contrary to predictions, such as positive effects of summer water availability in wetter parts of the range and positive responses to winter warming especially in milder areas. In general, we found wood frogs were more sensitive to changes in temperature or temperature interacting with precipitation than to changes in precipitation alone. Our results suggest that sensitivity to changes in climate cannot be predicted simply by knowing locations within the species’ climate envelope. Many climate processes did not affect population growth rates in the predicted direction based on range position. Processes such as species‐interactions, local adaptation, and interactions with the physical landscape likely affect the responses we observed. Our work highlights the need to measure demographic responses to changing climate.     

Water deficit in field-grown Gossypium hirsutum primarily limits net photosynthesis by decreasing stomatal conductance,increasing photorespiration,and increasing the ratio of dark respiration to gross photosynthesis

Much effort has been expended to improve irrigation efficiency and drought tolerance of agronomic crops; however, a clear understanding of the physiological mechanisms that interact to decrease source strength and drive yield loss has not been attained. To elucidate the underlying mechanisms contributing to inhibition of net carbon assimilation under drought stress, three cultivars of Gossypium hirsutum were grown in the field under contrasting irrigation regimes during the 2012 and 2013 growing season near Camilla, Georgia, USA. Physiological measurements were conducted on three sample dates during each growing season (providing a broad range of plant water status) and included, predawn and midday leaf water potential (Ψ_PD and Ψ_MD), gross and net photosynthesis, dark respiration, photorespiration, and chlorophyll a fluorescence. End-of-season lint yield was also determined. Ψ_PD ranged from −0.31 to −0.95 MPa, and Ψ_MD ranged from −1.02 to −2.67 MPa, depending upon irrigation regime and sample date. G. hirsutum responded to water deficit by decreasing stomatal conductance, increasing photorespiration, and increasing the ratio of dark respiration to gross photosynthesis, thereby limiting P_N and decreasing lint yield (lint yield declines observed during the 2012 growing season only). Conversely, even extreme water deficit, causing a 54% decline in P_N, did not negatively affect actual quantum yield, maximum quantum yield, or photosynthetic electron transport. It is concluded that P_N is primarily limited in drought-stressed G. hirsutum by decreased stomatal conductance, along with increases in respiratory and photorespiratory carbon losses, not inhibition or down-regulation of electron transport through photosystem II. It is further concluded that Ψ_PD is a reliable indicator of drought stress and the need for irrigation in field-grown cotton.