Thermal plasticity of photosynthesis: the role of acclimation in forest responses to a warming climate

The increasing air temperatures central to climate change predictions have the potential to alter forest ecosystem function and structure by exceeding temperatures optimal for carbon gain. Such changes are projected to threaten survival of sensitive species, leading to local extinctions, range migrations, and altered forest composition. This study investigated photosynthetic sensitivity to temperature and the potential for acclimation in relation to the climatic provenance of five species of deciduous trees, Liquidambar styraciflua, Quercus rubra, Quercus falcata, Betula alleghaniensis, and Populus grandidentata. Open‐top chambers supplied three levels of warming (+0, +2, and +4 °C above ambient) over 3 years, tracking natural temperature variability. Optimal temperature for CO₂ assimilation was strongly correlated with daytime temperature in all treatments, but assimilation rates at those optima were comparable. Adjustment of thermal optima was confirmed in all species, whether temperatures varied with season or treatment, and regardless of climate in the species' range or provenance of the plant material. Temperature optima from 17° to 34° were observed. Across species, acclimation potentials varied from 0.55 °C to 1.07 °C per degree change in daytime temperature. Responses to the temperature manipulation were not different from the seasonal acclimation observed in mature indigenous trees, suggesting that photosynthetic responses should not be modeled using static temperature functions, but should incorporate an adjustment to account for acclimation. The high degree of homeostasis observed indicates that direct impacts of climatic warming on forest productivity, species survival, and range limits may be less than predicted by existing models.     

Neglecting acclimation of photosynthesis under drought can cause significant errors in predicting leaf photosynthesis in wheat

Extreme climatic events, such as heat waves, cold snaps and drought spells, related to global climate change, have become more frequent and intense in recent years. Acclimation of plant physiological processes to changes in environmental conditions is a key component of plant adaptation to climate change. We assessed the temperature response of leaf photosynthetic parameters in wheat grown under contrasting water regimes and growth temperatures (T_growth). Two independent experiments were conducted under controlled conditions. In Experiment 1, two wheat genotypes were subjected to well-watered or drought-stressed treatments; in Experiment 2, the two water regimes combined with high, medium and low T_growth were imposed on one genotype. Parameters of a biochemical C₃-photosynthesis model were estimated at six leaf temperatures for each factor combination. Photosynthesis acclimated more to drought than to T_growth. Drought affected photosynthesis by lowering its optimum temperature (T_opt) and the values at T_opt of light-saturated net photosynthesis, stomatal conductance, mesophyll conductance, the maximum rate of electron transport (J_max) and the maximum rate of carboxylation by Rubisco (V_cmax). T_opt for V_cmax was up to 40°C under well-watered conditions but 24–34°C under drought. The decrease in photosynthesis under drought varied among T_growth but was similar between genotypes. The temperature response of photosynthetic quantum yield under drought was partly attributed to photorespiration but more to alternative electron transport. All these changes in biochemical parameters could not be fully explained by the changed leaf nitrogen content. Further model analysis showed that both diffusional and biochemical parameters of photosynthesis and their thermal sensitivity acclimate little to T_growth, but acclimate considerably to drought and the combination of drought and T_growth. The commonly used modelling approaches, which typically consider the response of diffusional parameters, but ignore acclimation responses of biochemical parameters to drought and T_growth, strongly overestimate leaf photosynthesis under variable temperature and drought.     

Response of plant functional traits of Leymus chinensis to extreme drought in Inner Mongolia grasslands

Plant Ecology - Understanding the effects of climate change, in particular, climate extremes on plant functional traits can provide a mechanistic basis for predicting how plant communities may be...     

A study of soil methane sink regulation in two grasslands exposed to drought and N fertilization

Oxidation by soil bacteria is the only biological sink for atmospheric methane (CH₄). There are substantial uncertainties regarding the global size of this sink, in part because the ecological controls of the involved processes are not well understood to date. We have investigated effects of severe summer drought and of nitrogen inputs (ammonium nitrate or cattle urine) on soil CH₄ fluxes in a field experiment. Soil moisture was the most important factor regulating the temporal dynamics of CH₄ fluxes. Simulated drought episodes altered the soil’s water balance throughout the year, increasing CH₄ oxidation by 50% on an annual basis. N fertilizers exerted only small and transient effects at the ecosystem level. Laboratory incubations suggested that effects differed between soil layers, with larger effects of drought and N application in the top soil than in deeper layers. With soil moisture being the primary controlling factor of methanotrophy, a detailed understanding of the ecosystem’s water balance is required to predict CH₄ budgets under future climatic conditions.     

Effects of a warmer climate on seed germination in the subarctic

Background and Aims

In a future warmer subarctic climate, the soil temperatures experienced by dispersed seeds are likely to increase during summer but may decrease during winter due to expected changes in snow depth, duration and quality. Because little is known about the dormancy-breaking and germination requirements of subarctic species, how warming may influence the timing and level of germination in these species was examined.

Methods

Under controlled conditions, how colder winter and warmer summer soil temperatures influenced germination was tested in 23 subarctic species. The cold stratification and warm incubation temperatures were derived from real soil temperature measurements in subarctic tundra and the temperatures were gradually changed over time to simulate different months of the year.

Key Results

Moderate summer warming (+2·5 °C) substantially accelerated germination in all but four species but did not affect germination percentages. Optimum germination temperatures (20/10°C) further decreased germination time and increased germination percentages in three species. Colder winter soil temperatures delayed the germination in ten species and decreased the germination percentage in four species, whereas the opposite was found in Silene acaulis. In most species, the combined effect of a reduced snow cover and summer warming resulted in earlier germination and thus a longer first growing season, which improves the chance of seedling survival. In particular the recruitment of (dwarf) shrubs (Vaccinium myrtillus, V. vitis-idaea, Betula nana), trees (Alnus incana, Betula pubescens) and grasses (Calamagrostis lapponica, C. purpurea) is likely to benefit from a warmer subarctic climate.

Conclusions

Seedling establishment is expected to improve in a future warmer subarctic climate, mainly by considerably earlier germination. The magnitudes of the responses are species-specific, which should be taken into account when modelling population growth and migration of subarctic species.Key words: Climate change, cold stratification, dwarf shrubs, germination percentage, incubation temperature, mean germination time, seedling establishment, seeds, Silene, subarctic species, Vaccinium, warming     

Low impact of climate change on subalpine grasslands in the Swiss Northern Alps

While phenological shifts and migration of isolated species under climate change have already been observed on alpine summits, very few studies have focused on community composition changes in subalpine grasslands. Here we use permanent plots monitored since 1954 and precisely located phytosociological censuses from 1970 to study compositional changes of subalpine grasslands in two distinct regions of the Swiss Northern Alps. In both areas, warming trends during the monitoring period were associated with changes in land management (abandonment of goat and sheep pasturing or grazing replaced by mowing). Old and recent inventories were compared with correspondence analyses (CA). Ecological indicator values, community‐affinities and biological traits of the species were used to infer the factors responsible for triggering the observed changes. In both regions, subalpine grasslands were stable with smaller changes than have previously been observed in alpine environments. Only a few species appeared or disappeared and changes were generally limited to increasing or decreasing frequency and cover of certain taxa. At one site, grazing abandonment favored fallow species. Some of these species were located at their upper altitudinal distribution limits and may have spread because of rising temperatures. In both areas, declining species were predominantly alpine and low‐growing species; their decline was probably due to increased competition (e.g., shadow) with more vigorous subalpine taxa no longer limited by grazing. We conclude that vegetation communities can respond rapidly to warming as long as colonization is facilitated by available space or structural change. In the subalpine grasslands studies, changes were mainly driven by land management. These communities have a dense vegetation cover and newly arriving herbaceous species preferring warmer conditions may take some time to establish themselves. However, climate disturbances, such as exceptional drought, may accelerate community changes by opening gaps for new species.     

Physiological acclimation to drought stress in Solidago canadensis

Solidago canadensis is an invasive species from North America that is spreading across Europe, Australia and temperate Asia. We hypothesized that the species' wide ecological amplitude is also based on its potential in hydraulic acclimation, and analyzed hydraulic and anatomical properties along a transect with decreasing soil humidity. Stem hydraulic conductivity, vulnerability to drought‐induced embolism, stomatal closure during dehydration and xylem‐anatomical parameters were quantified at three sites. At the humid site, specific hydraulic conductivity of stems (1.0 ± 0.2 kg m^–1 MPa^–1 s^–1) was about twofold higher, and leaf‐specific conductivity about 1.5 times higher (3.1 ± 0.5 kg m^–1 MPa^–1 s^–1) than at the dry site. Water potential (Ψ) at 50% loss of conductivity was ?3.7 ± 0.1 MPa at the dry site and ?3.1 ± 0.2 MPa at the humid site (September). Vulnerability to drought‐induced embolism decreased along the transect and over the vegetation period. At drier sites, stomata started closing at lower Ψ while complete stomatal closure was reached at less negative Ψ (12% of maximum stomatal conductance: –2.5 ± 0.0 and ?3.0 ± 0.2 MPa at the dry and humid site). The safety margin between stomatal closure and 50% loss of conductivity was 1.2 and 0.2 MPa at the dry and humid sites. The observed variability indicated an efficient acclimation in hydraulic conductivity and safety: plants at dry sites exhibited lower specific hydraulic conductivity, higher embolism resistance and broader safety margins, signifying a trade‐off between the hydraulic safety and efficiency. The observed intraspecific plasticity in hydraulic and anatomical traits may help to explain the invasive potential of this species.     

Multiscale topoedaphic heterogeneity increases resilience and resistance of a dominant grassland species to extreme drought and climate change

It is argued that the inclusion of spatially heterogeneous environments in biodiversity reserves will be an effective means of encouraging ecosystem resilience and plant community conservation under climate change. However, the resilience and resistance of plant populations to global change, the specific life‐history traits involved and the spatial scale at which environmentally driven demographic variation is expressed remains largely unknown for most plant groups. Here we address these questions by reporting an empirical investigation into the impacts of an unprecedented 3‐year drought on the demography, population growth rates (λ) and biogeographical distribution of core populations of the perennial grassland species Austrostipa aristiglumis in semiarid Australia. We use life‐history analysis and periodic matrix population models to specifically test the hypothesis that patch‐ and habitat‐scale variation in vital life‐history parameters result in spatial differences in the resilience and resistance of A. aristiglumis populations to extreme drought. We show that the development of critical soil water deficits during drought resulted in collapse of adult A. aristiglumis populations (λ?1), rapid interhabitat phytosociological change and overall contraction towards mesic refugia where populations were both more resistant and resilient to perturbation. Population models, combined with climatic niche analysis, suggest that, even in core areas, a significant reduction in size and habitat range of A. aristiglumis populations is likely under climate change expected this century. Remarkably, however, we show that even minor topographic variation (0.2–3 m) can generate significant variation in demographic parameters that confer population‐level resilience and resistance to drought. Our findings support the hypothesis that extreme climatic events have the capacity to induce rapid, landscape‐level shifts in core plant populations, but that the protection of topographically heterogeneous environments, even at small spatial scales, may play a key role in conserving biodiversity under climate change in the coming century.     

Contrasting thermal acclimation of leaf dark respiration and photosynthesis of Antarctic vascular plant species exposed to nocturnal warming

Leaf respiration and photosynthesis will respond differently to an increase in temperature during night, which can be more relevant in sensitive ecosystems such as Antarctica. We postulate that the plant species able to colonize the Antarctic Peninsula – Colobanthus quitensis (Kunth) Bartl. and Deschampsia antarctica Desv. – are able to acclimate their foliar respiration and to maintain photosynthesis under nocturnal warming to sustain a positive foliar carbon balance. We conducted a laboratory experiment to evaluate the effect of time of day (day and night) and nocturnal warming on dark respiration. Short (E₀ and Q₁₀) and long‐term acclimation of respiration, leaf carbohydrates, photosynthesis (A_sat) and foliar carbon balance (R/A) were evaluated. The results suggest that the two species have differential thermal acclimation respiration, where D. antarctica showed more thermosensitivity to short‐term changes in temperature than C. quitensis. Experimental nocturnal warming affected respiration at daytime differentially between the two species, with a significant increase of R₁₀ and A_sat in D. antarctica, while no changes on respiration were observed in C. quitensis. Long thermal treatments of the plants indicated that nocturnal but not diurnal respiration could acclimate in both species, and to a greater extent in C. quitensis. Non‐structural carbohydrates were related with respiration in C. quitensis but not in D. antarctica, suggesting that respiration in the former species is likely controlled by total soluble sugars and starch during day and night, respectively. Finally, foliar carbon balance was differentially improved under warming conditions in Antarctic plants by different mechanisms, with C. quitensis deploying respiratory acclimation, while D. antarctica increased its A_sat.     

Antioxidant response of wheat roots to drought acclimation

Wheat (Triticum aestivum L.) seedlings of a drought-resistant cv. C306 were subjected to severe water deficit directly or through stress cycles of increasing intensity with intermittent recovery periods. The antioxidant defense in terms of redox metabolites and enzymes in root cells and mitochondria was examined in relation to membrane damage. Acclimated seedlings exhibited higher relative water content and were able to limit the accumulation of H₂O₂ and membrane damage during subsequent severe water stress conditions. This was due to systematic up-regulation of superoxide dismutase, ascorbate peroxidase (APX), catalase, peroxidases, and ascorbate–glutathione cycle components at both the whole cell level as well as in mitochondria. In contrast, direct exposure of severe water stress to non-acclimated seedlings caused greater water loss, excessive accumulation of H₂O₂ followed by elevated lipid peroxidation due to the poor antioxidant enzyme response particularly of APX, monodehydroascorbate reductase, dehydroascorbate reductase, glutathione reductase, and ascorbate–glutathione redox balance. Mitochondrial antioxidant defense was found to be better than the cellular defense in non-acclimated roots. Termination of stress followed by rewatering leads to a rapid enhancement in all the antioxidant defense components in non-acclimated roots, which suggested that the excess levels of H₂O₂ during severe water stress conditions might have inhibited or down-regulated the antioxidant enzymes. Hence, drought acclimation conferred enhanced tolerance toward oxidative stress in the root tissue of wheat seedlings due to both reactive oxygen species restriction and well-coordinated induction of antioxidant defense.     

Zooplankton response to extreme drought in a large subtropical lake

Plankton data from 1997 to 2005 were used to examine impacts of a managed draw-down, subsequent drought and resulting historic low water levels (during 2000 and 2001) on the zooplankton of Lake Okeechobee, Florida. Prior to the drought the lake supported less than 150 ha of submerged vegetation. Following the drought, over 15,000 ha of submerged vegetation developed around the lake shore and conditions favored greater survival of age 0 fish. The zooplankton changed significantly from the pre- to post-drought period, including: (a) a near-complete loss of all dominant species of cladocerans and rotifers; and (b) an abrupt transition to a community with over 80% of total biomass comprised of Arctodiaptomus dorsalis, a calanoid copepod previously described as being resistant to fish predation. These changes persisted over a 5 year post-drought sampling period. In contrast, there were no systematic changes in biomass of bacteria, phytoplankton, inedible cyanobacteria, algal cell size, suspended solids, or any other physical or chemical attributes known to affect zooplankton in shallow lakes. Evidence points towards increased predation by fish, and perhaps invertebrates, as factors responsible for loss of cladocerans and rotifers following the drought, and indicates a need for future research to link changes in water level to shifts in predation pressure in this and other shallow lakes. Handling editor: S.I. Dodson     

Photochemical response to drought acclimation in two sunflower genotypes

The effects of drought acclimation on CO₂ assimilation and light utilization were investigated in two sunflower genotypes ( Helianthus annuus L., T32 and Viki) in relation to water deficit and/or high light conditions. Drought interaction with PSII efficiency was observed in the genotype T32 with a sustained decrease in the potential photochemical efficiency of PSII, F_n/F_m. In response to drought acclimation, T32 displayed some tendency to accumulate closed PSII traps (higher value of 1-qp) without an enhancement of thermal deactivation (Stem-Volmer non-photochemical quenching, NPQ). Irrespective of the growth conditions (growth chamber or greenhouse), only Viki was responsive to drought acclimation, with (1) increased net photosynthesis in well-watered plants, (2) higher maintenance of photochemical electron transfer under water deficit and/or high light, (3) limited PSII inactivation (lower value of 1-qp) through increased non-photochemical energy dissipation (Stern-Volmer NPQ) which was readily reversible even at low leaf water potentials, and (4) higher F_v/F_m recovery after high light treatment. Additionally, drought acclimation delayed turgor loss during subsequent water stress in Viki. Thus, the response to drought acclimation, with an adjustment of water relations and of energy utilization by PSII, was observed under both growth conditions and was mainly genotype dependent.     

Unexpected patterns of sensitivity to drought in three semi-arid grasslands

Global climate models forecast an increase in the frequency and intensity of extreme weather events, including severe droughts. Based on multi-year relationships between precipitation amount and aboveground annual net primary production (ANPP), semi-arid grasslands are projected to be among the most sensitive ecosystems to changes in precipitation. To assess sensitivity to drought, as well as variability within the shortgrass steppe biome, we imposed moderate and severe rainfall reductions for two growing seasons in three undisturbed grasslands that varied in soil type and climate. We predicted strong drought-induced reductions in ANPP at all sites and greater sensitivity to drought in sites with lower average precipitation, consistent with continental-scale patterns. Identical experimental infrastructure at each site reduced growing season rainfall events by 50 or 80%, and significantly reduced average soil moisture in both years (by 21 and 46% of control levels, respectively). Despite reductions in soil moisture, ANPP responses varied unexpectedly-from no reduction in ANPP to a 51% decrease. Although sensitivity to drought was highest in the semi-arid grassland with lowest mean annual precipitation, patterns in responses to drought across these grasslands were also strongly related to rainfall event size. When growing season rainfall patterns were dominated by many smaller events, ANPP was significantly reduced by drought but not when rainfall patterns were characterized by large rain events. This interaction between drought sensitivity and rainfall event size suggests that ANPP responses to future droughts may be reduced if growing season rainfall regimes also become more extreme.     

Temperature acclimation of net photosynthesis in relation to growth of a cold hardy Chrysanthemum

Summary Direct and interactive effects of daylength and temperature preconditioning of above- and below-ground parts of Chrysanthemum morifolium Ramat. Astrid are examined in relation to net photosynthesis. Most previous work on thermal preconditioning has concentrated on the effects of a given thermal regime on entire plants, without attempts to separate the effects of temperature acting on shoots vs. below-ground organs (roots and rhizomes).Vegetatively propagated cuttings were placed under 2 different photoperiods and 4 different thermal regimes for a total of 8 treatments. Subsequent to 40-day preconditioning regimes, the temperature-dependence of net photosynthesis was determined between 8 and 40 degrees C under a photon flux density of about 540 E·m^-2·s^-1. The photosynthetic rate data were treated by multiple analysis of variance to examine direct and interactive factor effects.Each of the preconditioning factors (daylength, shoot preconditioning temperature, root preconditioning temperature) individually affected the measured photosynthesis rates, and the effects were very highly significant (p<0.001). In the two-way interactive effects, both daylength X shoot environment, and shoot environment X root environment were very highly significant (p<0.001) and daylength X root environment was significant (p<0.05). Comparison of sums of squares indicated the greatest effect was due to shoot preconditioning environment, while the least important was daylength. Photosynthesis temperature at the time of gas exchange measurements did not interact with the preconditioning temperatures or with daylength, with the implication that preconditioning affected the elevation (absolute rates) of the photosynthesis curves but not the shapes of the curves.The data are further discussed in relation to models of resource allocation and theory of natural selection for optimization.     

Ecophysiology of a wild platyopuntia exposed to prolonged drought

The effects of reduced annual rainfall over a 7-year period on the daily patterns of gas exchange were analyzed for a wild platyopuntia (Opuntia robusta) growing in rocky soil. In spite of the prolonged water stress, stomata opened at night and net carbon gain was observed for eight 1-day sampling periods during 2000. Daily net CO₂ uptake increased more than five-fold from the end of the dry spring season to the middle of the summer wet season. Curtailment in the formation of both new cladodes and new roots is considered to be an avoidance strategy to water scarcity, which in combination with net CO₂ uptake responses to temperature and relative humidity at night, and perhaps association with arbuscular-mycorrhizal fungi, were the most important adaptations for O. robusta to the prolonged drought conditions that prevailed in its rocky environment. Such adaptations of O. robusta are in addition to the classical physiological and structural modifications of CAM plants to tolerate prolonged drought.     

1982-2015年中国温带不同草地植被枯黄期对极端气候事件的响应

随着极端气候事件频率和强度的增加,植被物候正在发生深刻的变化。然而,植被枯黄期(EGS)对极端气候的响应机制目前尚未厘清,特别是对于干旱半干旱地区的草地而言。因此,聚焦我国温带草地,基于1982—2015年全球监测与模型研究工作组归一化植被指数(GIMMS NDVI3g)长时间序列数据提取草地物候参数,并分析其时空变化规律;运用随机森林模型等方法探究温带草地EGS对极端气候变化的响应特征。结果表明：(1)全区多年平均EGS主要发生于270—290儒略日(DOY),59.8%的区域呈延迟趋势,其中显著延迟(P<0.05)的区域分布在新疆天山、阿尔泰山一带和准噶尔盆地西部、黄土高原北部、呼伦贝尔高原的西部和东北小兴安岭。(2)EGS与极端气温暖极值(日最低气温的最大值、日最高气温的最大值、暖夜日数、暖昼日数)之间均以广泛的正相关关系为主;相比之下,极端降水事件与EGS之间的关系相对比较复杂,这与各草地类型自身的生理策略和所处环境密切相关。(3)整体而言,持续干旱日数、气温日较差和暖夜日数对全域草地EGS动态变化具有极大的重要性。就不同草地类型而言,温带草甸草原主要受到气温日较差的影响...     

Water use efficiency,photosynthesis, extreme temperature and drought tolerance of desert plant seedlings

The seedlings of twenty eight annual and perennial desert plants belonging to fifteen families were raised under controlled growth conditions and subjected to a range of tests. The water use efficiency, photosynthesis, extreme temperatures and drought tolerance were monitored. The results were discussed for their ecological significance as a basic information to applied research. Seedlings which were more tolerant to extreme high and low temperatures and drought were more capable of controlling water loss through better water use efficiency. At temperatures below 25°C, the annual plants had higher photosynthetic rate than perennials, whereas at higher temperatures, the perennials were advantaged.     

Strategies for acclimation to seasonal drought in Ceratonia siliqua leaves

Plants of Ceratonia siliqua L. (carob tree) were subjected to a slow cycle of soil water depletion in summer and in winter. Recently mature and fully turgid leaves were squeezed in a pressure chamber in order to analyse changes in the components of water potential related to the acclimation to drought. Mathematical expressions were fitted to the primary data in order to calculate the elasticity modulus and the dependence of turgor pressure on relative osmotic water content. Osmotic and water potentials decreased significantly in leaves acclimated during the summer (about 0.7 and 0.9 MPa decrease, respectively) whereas in winter the elastic properties of the wall had changed: modulus of the non-stressed leaves was 26 MPa compared to 7.5 MPa in leaves subjected to drought stress. The results indicate that Ceratonia leaves can, to some extent, maintain turgor under situations of soil drought, using different strategies according to the season.     

Stimulation of both photosynthesis and respiration in response to warmer and drier conditions in a boreal peatland ecosystem

Peatland ecosystems have been consistent carbon (C) sinks for millennia, but it has been predicted that exposure to warmer temperatures and drier conditions associated with climate change will shift the balance between ecosystem photosynthesis and respiration providing a positive feedback to atmospheric CO₂ concentration. Our main objective was to determine the sensitivity of ecosystem photosynthesis, respiration and net ecosystem production (NEP) measured by eddy covariance, to variation in temperature and water table depth associated with interannual shifts in weather during 2004–2009. Our study was conducted in a moderately rich treed fen, the most abundant peatland type in western Canada, in a region (northern Alberta) where peatland ecosystems are a significant landscape component. During the study, the average growing season (May–October) water depth declined approximately 38 cm, and temperature [expressed as cumulative growing degree days (GDD, March–October)] varied approximately 370 GDD. Contrary to previous predictions, both ecosystem photosynthesis and respiration showed similar increases in response to warmer and drier conditions. The ecosystem remained a strong net sink for CO₂ with an average NEP (± SD) of 189 ± 47 g C m^?2 yr^?1. The current net CO₂ uptake rates were much higher than C accumulation in peat determined from analyses of the relationship between peat age and cumulative C stock. The balance between C addition to, and total loss from, the top 0–30 cm depth (peat age range 0–70 years) of shallow peat cores averaged 43 ± 12 g C m^?2 yr^?1. The apparent long‐term average rate of net C accumulation in basal peat samples was 19–24 g C m^?2 yr^?1. The difference between current rates of net C uptake and historical rates of peat accumulation is likely a result of vegetation succession and recent increases in tree establishment and productivity.