Elucidating the impact of temperature variability and extremes on cereal croplands through remote sensing

Remote sensing‐derived wheat crop yield‐climate models were developed to highlight the impact of temperature variation during thermo‐sensitive periods (anthesis and grain‐filling; TSP) of wheat crop development. Specific questions addressed are: can the impact of temperature variation occurring during the TSP on wheat crop yield be detected using remote sensing data and what is the impact? Do crop critical temperature thresholds during TSP exist in real world cropping landscapes? These questions are tested in one of the world's major wheat breadbaskets of Punjab and Haryana, north‐west India. Warming average minimum temperatures during the TSP had a greater negative impact on wheat crop yield than warming maximum temperatures. Warming minimum and maximum temperatures during the TSP explain a greater amount of variation in wheat crop yield than average growing season temperature. In complex real world cereal croplands there was a variable yield response to critical temperature threshold exceedance, specifically a more pronounced negative impact on wheat yield with increased warming events above 35 °C. The negative impact of warming increases with a later start‐of‐season suggesting earlier sowing can reduce wheat crop exposure harmful temperatures. However, even earlier sown wheat experienced temperature‐induced yield losses, which, when viewed in the context of projected warming up to 2100 indicates adaptive responses should focus on increasing wheat tolerance to heat. This study shows it is possible to capture the impacts of temperature variation during the TSP on wheat crop yield in real world cropping landscapes using remote sensing data; this has important implications for monitoring the impact of climate change, variation and heat extremes on wheat croplands.     

Post‐flowering night respiration and altered sink activity account for high night temperature‐induced grain yield and quality loss in rice (Oryza sativa L.)

High night temperature (HNT) is a major constraint to sustaining global rice production under future climate. Physiological and biochemical mechanisms were elucidated for HNT‐induced grain yield and quality loss in rice. Contrasting rice cultivars (N22, tolerant; Gharib, susceptible; IR64, high yielding with superior grain quality) were tested under control (23°C) and HNT (29°C) using unique field‐based tents from panicle initiation till physiological maturity. HNT affected 1000 grain weight, grain yield, grain chalk and amylose content in Gharib and IR64. HNT increased night respiration (Rn) accounted for higher carbon losses during post‐flowering phase. Gharib and IR64 recorded 16 and 9% yield reduction with a 63 and 35% increase in average post‐flowering Rn under HNT, respectively. HNT altered sugar accumulation in the rachis and spikelets across the cultivars with Gharib and IR64 recording higher sugar accumulation in the rachis. HNT reduced panicle starch content in Gharib (22%) and IR64 (11%) at physiological maturity, but not in the tolerant N22. At the enzymatic level, HNT reduced sink strength with lower cell wall invertase and sucrose synthase activity in Gharib and IR64, which affected starch accumulation in the developing grain, thereby reducing grain weight and quality. Interestingly, N22 recorded lower Rn‐mediated carbon losses and minimum impact on sink strength under HNT. Mechanistic responses identified will facilitate crop models to precisely estimate HNT‐induced damage under future warming scenarios.     

Body size response to warming: time of the season matters in a tephritid fly

Whether shrinking body size is a universal response to climate change remains controversial. Moreover, the mechanisms underlying body size shifts are poorly understood. Here, assuming that life history traits evolve to maximize fitness according to life history plasticity theory, we hypothesized that under global warming temperate multivoltine insects should emerge earlier with a smaller body mass in the early growing season, but emerge later with a larger body mass in the late season. We tested this hypothesis by conducting two field artificial warming experiments in an alpine meadow: 1) with one pre‐dispersal seed predator species (tephritid flies, Tephritis femoralis) and its two host‐plant species flowering in early and late growing seasons, respectively, and 2) with the tephritid flies and one host species with a flowering season that occupies parts of both the early and late growing seasons. These experiments were complemented by a microcosm chamber warming experiment, in which increasing and decreasing temperature trends were set to simulate temperature variation pattern in early and late growing seasons, respectively, but photoperiod was held constant. Warming generally advanced the adult emergence and decreased the body size (adult body mass) in the early season but delayed the emergence and increased the size in the late season in both field experiments, indicating that the seasonally different effects of warming on the fly body size was constant regardless of host‐plant identity. The chamber warming resulted in consistent responses of emerging timing and body size to the simulated seasonal warming, demonstrating that the temperature increase per se and its interaction with direction of temperature change, but not other correlated effects, should be primarily responsible for the observed contrasting shifts of body size at different times of the season. Our results indicate that taking into account temperate seasonal patterns of temperature variation could be of general importance for predicting animal body size changes in the warmed future.     

Phenological Sensitivity of Early and Late Flowering Species Under Seasonal Warming and Altered Precipitation in a Seminatural Temperate Grassland Ecosystem

Shifts in flowering phenology of plants are indicators of climate change. The great majority of existing phenological studies refer solely to gradual warming. However, knowledge on how flowering phenology responds to changes in seasonal variation of warming and precipitation regimes is missing. We report the onset of 22 early (flowering before/within May) and 23 late flowering (flowering after May) species in response to manipulated seasonal warming (equal to + 1.2°C; last 100-year summer/winter warming), additional winter rainfall, and modified precipitation variability (including a 1000-year extreme drought event followed by heavy rainfall) over the growing season in two consecutive years for a species-rich temperate grassland ecosystem. The average onset of flowering (over 2 years) was significantly advanced 3.1 days by winter warming and 1.5 days by summer warming compared to control. Early flowering species responded to seasonal warming in both years, while late-flowering species responded in only 1 year to summer warming. The average onset of early flowering species was significantly advanced, 4.9 days by winter warming and 2.3 days by summer warming. Species-specific analysis showed that even within the early flowering community there were divergences. A positive correlation between plant height and shift in flowering onset was detected under winter warming (R² = 0.20, p = 0.005). The average onsets of early and late flowering community were affected by neither winter rain nor growing season precipitation variability. Seasonal differences in warming, and particularly winter warming, might alter community dynamics among early and late flowering species which can cause shifts in the seasonal performances of temperate ecosystems.     

The impact of temperature variability on wheat yields

With current annual production at over 600 million tonnes, wheat is the third largest crop in the world behind corn and rice, and an essential source of carbohydrates for millions of people. While wheat is grown over a wide range of environments, it is common in the major wheat‐producing countries for grain filling to occur when soil moisture is declining and temperature is increasing. Average global temperatures have increased over the last decades and are predicted to continue rising, along with a greater frequency of extremely hot days. Such events have already been reported for major wheat growing regions in the world. However, the direct impact of past temperature variability and changes in averages and extremes on wheat production has not been quantified. Attributing changes in observed yields over recent decades to a single factor such as temperature is not possible due to the confounding effects of other factors. By using simulation modelling, we were able to separate the impact of temperature from other factors and show that the effect of temperature on wheat production has been underestimated. Surprisingly, observed variations in average growing‐season temperatures of ±2 °C in the main wheat growing regions of Australia can cause reductions in grain production of up to 50%. Most of this can be attributed to increased leaf senescence as a result of temperatures >34 °C. Temperature conditions during grain filling in the major wheat growing regions of the world are similar to the Australian conditions during grain filling. With average temperatures and the frequency of heat events projected to increase world‐wide with global warming, yield reductions due to higher temperatures during the important grain‐filling stage alone could substantially undermine future global food security. Adaptation strategies need to be considered now to prevent substantial yield losses in wheat from increasing future heat stress.     

Climate Change: Implications for the Yield of Edible Rice

Global warming affects not only rice yield but also grain quality. A better understanding of the effects of climate factors on rice quality provides information for new breeding strategies to develop varieties of rice adapted to a changing world. Chalkiness is a key trait of physical quality, and along with head rice yield, is used to determine the price of rice in all markets. In the present study, we show that for every ∼1% decrease in chalkiness, an increase of ∼1% in head rice yield follows, illustrating the dual impact of chalk on amount of marketable rice and its value. Previous studies in controlled growing conditions report that chalkiness is associated with high temperature. From 1980–2009 at IRRI, Los Baños, the Philippines, annual minimum and mean temperatures, and diurnal variation changed significantly. The objective of this study was to determine how climate impacts chalkiness in field conditions over four wet and dry seasons. We show that low relative humidity and a high vapour pressure deficit in the dry season associate with low chalk and high head rice yield in spite of higher maximum temperature, but in the opposite conditions of the wet season, chalk is high and head rice yield is low. The data therefore suggest that transpirational cooling is a key factor affecting chalkiness and head rice yield, and global warming per se might not be the major factor that decreases the amount and quality of rice, but other climate factors in combination, that enable the crop to maintain a cool canopy.     

Diel and seasonal variations in the thermal biology of San Cristobal Lava Lizards (Microlophus bivittatus)

Thermal biology, and therefore energy acquisition and survival, of ectotherms can be affected by diel and seasonal patterns of environmental temperatures. Galápagos Lava Lizards live in seasonal environments that are characterized by a warm and wet period when reproductive activity is maximal, and cooler and drier period. With the use of radiotelemetric techniques to record lizard surface temperatures (T_s), we studied the thermal ecology of the San Cristóbal Lava Lizard (Microlophus bivittatus) during both the warm and cool seasons over two years. During the diel activity period and when operative temperatures exceeded T_set-min, at least on rock faces without canopy, 52% or less of the T_s observations fell within the laboratory-determined T_set range (36–40 °C). Therefore, lizards may have avoided very warm midday temperatures in shaded microhabitats and the lag times in changes in T_s values occurred as operative temperatures rose rapidly during late morning warming phase. Lizards effectively thermoregulated during a year with moderate warm season temperatures and during a cool season that was unseasonably warm. In contrast, lizards less effectively thermoregulated during the warmest and coolest years of the study. We did not detect intersexual differences in thermoregulation although males may thermoregulate less effectively than do females during the cool season although we were unable to detect significant differences using our nonparametric statistical techniques.     

Deterministic Plio-Pleistocene extinctions in the European cool-temperate tree flora

Extinctions have been important in the shaping of modern phytogeographic patterns. A classic example is the heavy Plio‐Pleistocene losses that have caused Europe to have a depauperate temperate tree flora compared to eastern North America and eastern Asia. To investigate the mechanisms involved in this extinction event, I test the hypothesis that the present European status (extinct, relictual, or widespread) of cool‐temperate tree genera found in Pliocene Europe is predictable from their modern climatic requirements. As a prerequisite for this analysis, I test for genus‐level conservatism in climatic requirements by comparing congeneric values across Europe, eastern Asia and North America, and find strong evidence hereof. I find a high degree of ecological determinism in the fate of European Pliocene tree genera, still widespread taxa being more tolerant of cold growing season and winter temperatures than extinct and relictual taxa, and relictual taxa being more drought tolerant than extinct taxa.     

A natural heating experiment: Phenotypic and genotypic responses of plant phenology to geothermal soil warming

Under global warming, the survival of many populations of sedentary organisms in seasonal environments will largely depend on their ability to cope with warming in situ by means of phenotypic plasticity or adaptive evolution. This is particularly true in high‐latitude environments, where current growing seasons are short, and expected temperature increases large. In such short‐growing season environments, the timing of growth and reproduction is critical to survival. Here, we use the unique setting provided by a natural geothermal soil warming gradient (Hengill geothermal area, Iceland) to study the response of Cerastium fontanum flowering phenology to temperature. We hypothesized that trait expression and phenotypic selection on flowering phenology are related to soil temperature, and tested the hypothesis that temperature‐driven differences in selection on phenology have resulted in genetic differentiation using a common garden experiment. In the field, phenology was related to soil temperature, with plants in warmer microsites flowering earlier than plants at colder microsites. In the common garden, plants responded to spring warming in a counter‐gradient fashion; plants originating from warmer microsites flowered relatively later than those originating from colder microsites. A likely explanation for this pattern is that plants from colder microsites have been selected to compensate for the shorter growing season by starting development at lower temperatures. However, in our study we did not find evidence of variation in phenotypic selection on phenology in relation to temperature, but selection consistently favoured early flowering. Our results show that soil temperature influences trait expression and suggest the existence of genetically based variation in flowering phenology leading to counter‐gradient local adaptation along a gradient of soil temperatures. An important implication of our results is that observed phenotypic responses of phenology to global warming might often be a combination of short‐term plastic responses and long‐term evolutionary responses, acting in different directions.     

The responses of microbial temperature relationships to seasonal change and winter warming in a temperate grassland

Microorganisms dominate the decomposition of organic matter and their activities are strongly influenced by temperature. As the carbon (C) flux from soil to the atmosphere due to microbial activity is substantial, understanding temperature relationships of microbial processes is critical. It has been shown that microbial temperature relationships in soil correlate with the climate, and microorganisms in field experiments become more warm‐tolerant in response to chronic warming. It is also known that microbial temperature relationships reflect the seasons in aquatic ecosystems, but to date this has not been investigated in soil. Although climate change predictions suggest that temperatures will be mostly affected during winter in temperate ecosystems, no assessments exist of the responses of microbial temperature relationships to winter warming. We investigated the responses of the temperature relationships of bacterial growth, fungal growth, and respiration in a temperate grassland to seasonal change, and to 2 years’ winter warming. The warming treatments increased winter soil temperatures by 5–6°C, corresponding to 3°C warming of the mean annual temperature. Microbial temperature relationships and temperature sensitivities (Q₁₀) could be accurately established, but did not respond to winter warming or to seasonal temperature change, despite significant shifts in the microbial community structure. The lack of response to winter warming that we demonstrate, and the strong response to chronic warming treatments previously shown, together suggest that it is the peak annual soil temperature that influences the microbial temperature relationships, and that temperatures during colder seasons will have little impact. Thus, mean annual temperatures are poor predictors for microbial temperature relationships. Instead, the intensity of summer heat‐spells in temperate systems is likely to shape the microbial temperature relationships that govern the soil‐atmosphere C exchange.     

Independent effects of warming and nitrogen addition on plant phenology in the Inner Mongolian steppe

Background and Aims

Phenology is one of most sensitive traits of plants in response to regional climate warming. Better understanding of the interactive effects between warming and other environmental change factors, such as increasing atmosphere nitrogen (N) deposition, is critical for projection of future plant phenology.

Methods

A 4-year field experiment manipulating temperature and N has been conducted in a temperate steppe in northern China. Phenology, including flowering and fruiting date as well as reproductive duration, of eight plant species was monitored and calculated from 2006 to 2009.

Key Results

Across all the species and years, warming significantly advanced flowering and fruiting time by 0·64 and 0·72 d per season, respectively, which were mainly driven by the earliest species (Potentilla acaulis). Although N addition showed no impact on phenological times across the eight species, it significantly delayed flowering time of Heteropappus altaicus and fruiting time of Agropyron cristatum. The responses of flowering and fruiting times to warming or N addition are coupled, leading to no response of reproductive duration to warming or N addition for most species. Warming shortened reproductive duration of Potentilla bifurca but extended that of Allium bidentatum, whereas N addition shortened that of A. bidentatum. No interactive effect between warming and N addition was found on any phenological event. Such additive effects could be ascribed to the species-specific responses of plant phenology to warming and N addition.

Conclusions

The results suggest that the warming response of plant phenology is larger in earlier than later flowering species in temperate grassland systems. The effects of warming and N addition on plant phenology are independent of each other. These findings can help to better understand and predict the response of plant phenology to climate warming concurrent with other global change driving factors.     

Association between rainfall seasonality and the flowering of epiphytic plants in a Neotropical montane forest

The association between the reproductive phenology of epiphytic communities with environmental and ecological factors remains largely unexplored. Because epiphytes depend on environmental moisture, seasonal changes in moisture conditions likely act as the primary determinants of their reproductive timing. We examined whether water limitation or pollinator competition structures the flowering phenologies of an epiphytic community in a seasonal mountain forest in Costa Rica. Additionally, we addressed the environmental factors that might trigger floral induction. Using a 24‐month dataset of bimonthly flowering records from 104 species, we found high seasonality of flowering at the species level but somewhat lower seasonality at the community level. The flowering mid‐dates of most epiphytes, particularly from monocotyledonous species, occurred during the wettest months, as predicted if water limitation structures flowering. The increased moisture and nutrient availability during the rainy season give epiphytes the resources needed to complete floral development and anthesis, and later fruit and seed maturation. The observed flowering pattern of epiphytes coincides with reproductive patterns of terrestrial herbs and shrubs from seasonal tropical ecosystems, and suggests shared constraints to sexual reproduction in both ecological guilds under similar climatic conditions. In contrast, flowering patterns of congeneric epiphytes in the same pollination guild mostly did not follow the expectations of a pollinator competition scenario. Finally, we discuss the possible combined effect of precipitation, temperature, and daily insolation on floral induction of epiphytic plants.     

Temperature alone does not explain phenological variation of diverse temperate plants under experimental warming

Anthropogenic climate change has altered temperate forest phenology, but how these trends will play out in the future is controversial. We measured the effect of experimental warming of 0.6–5.0 °C on the phenology of a diverse suite of 11 plant species in the deciduous forest understory (Duke Forest, North Carolina, USA) in a relatively warm year (2011) and a colder year (2013). Our primary goal was to dissect how temperature affects timing of spring budburst, flowering, and autumn leaf coloring for functional groups with different growth habits, phenological niches, and xylem anatomy. Warming advanced budburst of six deciduous woody species by 5–15 days and delayed leaf coloring by 18–21 days, resulting in an extension of the growing season by as much as 20–29 days. Spring temperature accumulation was strongly correlated with budburst date, but temperature alone cannot explain the diverse budburst responses observed among plant functional types. Ring‐porous trees showed a consistent temperature response pattern across years, suggesting these species are sensitive to photoperiod. Conversely, diffuse‐porous species responded differently between years, suggesting winter chilling may be more important in regulating budburst. Budburst of the ring‐porous Quercus alba responded nonlinearly to warming, suggesting evolutionary constraints may limit changes in phenology, and therefore productivity, in the future. Warming caused a divergence in flowering times among species in the forest community, resulting in a longer flowering season by 10‐16 days. Temperature was a good predictor of flowering for only four of the seven species studied here. Observations of interannual temperature variability overpredicted flowering responses in spring‐blooming species, relative to our warming experiment, and did not consistently predict even the direction of flowering shifts. Experiments that push temperatures beyond historic variation are indispensable for improving predictions of future changes in phenology.     

Impacts of extreme winter warming in the sub‐Arctic: growing season responses of dwarf shrub heathland

Climate change scenarios predict an increased frequency of extreme climatic events. In Arctic regions, one of the most profound of these are extreme and sudden winter warming events in which temperatures increase rapidly to above freezing, often causing snow melt across whole landscapes and exposure of ecosystems to warm temperatures. Following warming, vegetation and soils no longer insulated below snow are then exposed to rapidly returning extreme cold. Using a new experimental facility established in sub‐Arctic dwarf shrub heathland in northern Sweden, we simulated an extreme winter warming event in the field and report findings on growth, phenology and reproduction during the subsequent growing season. A 1‐week long extreme winter warming event was simulated in early March using infrared heating lamps run with or without soil warming cables. Both single short events delayed bud development of Vaccinium myrtillus by up to 3 weeks in the following spring (June) and reduced flower production by more than 80%: this also led to a near‐complete elimination of berry production in mid‐summer. Empetrum hermaphroditum also showed delayed bud development. In contrast, Vaccinium vitis‐idaea showed no delay in bud development, but instead appeared to produce a greater number of actively growing vegetative buds within plots warmed by heating lamps only. Again, there was evidence of reduced flowering and berry production in this species. While bud break was delayed, growing season measurements of growth and photosynthesis did not reveal a differential response in the warmed plants for any of the species. These results demonstrate that a single, short, extreme winter warming event can have considerable impact on bud production, phenology and reproductive effort of dominant plant species within sub‐Arctic dwarf shrub heathland. Furthermore, large interspecific differences in sensitivity are seen. These findings are of considerable concern, because they suggest that repeated events may potentially impact on the biodiversity and productivity of these systems should these extreme events increase in frequency as a result of global change. Although climate change may lengthen the growing season by earlier spring snow melt, there is a profound danger for these high‐latitude ecosystems if extreme, short‐lived warming in winter exposes plants to initial warm temperatures, but then extreme cold for the rest of the winter. Work is ongoing to determine the longer term and wider impacts of these events.     

Variation in the harvest index of tropical maize: evaluation of recent evidence from Mexico and Malawi

In temperate zones, the potential grain yield of wheat has increased during the twentieth century owing to progressive increases in the harvest index of new varieties, which are principally associated with reduction in plant stature. Crop biomass has not increased substantially. In contrast, the potential grain yield of maize in the USA has increased owing to progressive increases in biomass, principally associated with selection for grain yield at higher population density. Harvest index was already around 0.5 for recommended varieties in 1930, and has not increased significantly since. However, for both crops, the harvest index of a given variety has proved to be a highly‐heritable character, except under severe stress. Less is known about the physiology of tropical maize. This paper reviews evidence from Mexico and Malawi that tropical maize can respond to selection for reduced stature following the same pattern as temperate wheat, but, under other circumstances, the magnitude of harvest index is not highly heritable, varying inconsistently with season, management and environment. It is proposed that these differences arise out of the unique vulnerability of the grain‐setting process around flowering. The plasticity of harvest index under long, favourable conditions, however, remains to be explained, although it is probably also related to the events around grain setting. Nevertheless, to the subsistence farmer, higher harvest index may not be a high priority in crop improvement, because of the need for large quantities of high‐quality stover.     

Phenological changes in intertidal con‐specific gastropods in response to climate warming

There is substantial evidence from terrestrial and freshwater systems of species responding to climate change through changes in their phenology. In the marine environment, however, there is less evidence. Using historic (1946–1949) and contemporary (2003–2007) data, collected from rocky shores of south‐west Britain, we investigated the affect of recent climate warming on the reproductive phenology of two con‐specific intertidal limpet grazers, with cool/boreal and warm/lusitanian centres of distribution. Reproductive development in the southern limpet, Patella depressa, has advanced, on average, 10.2 days per decade since the 1940s, with a longer reproductive season and more of the population reproductively active. The peak in the proportion of the population in advanced stages of gonad development was positively correlated with sea surface temperature (SST) in late spring/early summer, which has increased between the 1940s and 2000s. The advance in peak reproductive development of this species is double the average observed for terrestrial and freshwater systems and indicates, along with other studies, that marine species may be responding faster to climate warming. In contrast, the northern limpet, Patella vulgata, has experienced a delay in the timing of its reproductive development (on average 3.3 days per decade), as well as an increase in reproductive failure years and a reduction in the proportion of the population reaching advanced gonad stages. These results are the first to demonstrate a delay in the reproductive development of a cool‐temperate, winter spawner, towards cooler more favourable environmental conditions in response to climate warming. Such a delay in spawning will potentially lead to trophic miss‐matches, resulting in a rapid nonlinear decline of this species.     

Physiological and biochemical characterization of NERICA‐L‐44: a novel source of heat tolerance at the vegetative and reproductive stages in rice

The predicted increase in the frequency and magnitude of extreme heat spikes under future climate can reduce rice yields significantly. Rice sensitivity to high temperatures during the reproductive stage is well documented while the same during the vegetative stage is more speculative. Hence, to identify and characterize novel heat‐tolerant donors for both the vegetative and reproductive stages, 71 rice accessions, including approximately 75% New Rice for Africa (NERICAs), were phenotyped across field experiments during summer seasons in Delhi, India, and in a controlled environment study at International Rice Research Institute , Philippines. NERICA‐L‐44 (NL‐44) recorded high seedling survival (52%) and superior growth and greater reproductive success exposed to 42.2°C (sd ± 2.3) under field conditions. NL‐44 and the heat‐tolerant check N22 consistently displayed lower membrane damage and higher antioxidant enzymes activity across leaves and spikelets. NL‐44 recorded 50–60% spikelet fertility, while N22 recorded 67–79% under controlled environment temperature of 38°C (sd ±1.17), although both had about 87% fertility under extremely hot field conditions. N22 and NL‐44, exposed to heat stress (38°C), had similar pollen germination percent and number of pollen tubes reaching the ovary. NL‐44 maintained low hydrogen peroxide production and non‐photochemical quenching (NPQ) with high photosynthesis while N22 avoided photosystem II damage through high NPQ under high‐temperature stress. NL‐44 with its reproductive stage resilience to extreme heat stress, better antioxidant scavenging ability in both vegetative tissue and spikelets and superior yield and grain quality is identified as a novel donor for increasing heat tolerance at both the vegetative and reproductive stages in rice.     

Yield response of field‐grown soybean exposed to heat waves under current and elevated [CO2]

Elevated atmospheric CO₂ concentration ([CO₂]) generally enhances C₃ plant productivity, whereas acute heat stress, which occurs during heat waves, generally elicits the opposite response. However, little is known about the interaction of these two variables, especially during key reproductive phases in important temperate food crops, such as soybean (Glycine max). Here, we grew soybean under elevated [CO₂] and imposed high‐ (+9°C) and low‐ (+5°C) intensity heat waves during key temperature‐sensitive reproductive stages (R1, flowering; R5, pod‐filling) to determine how elevated [CO₂] will interact with heat waves to influence soybean yield. High‐intensity heat waves, which resulted in canopy temperatures that exceeded optimal growth temperatures for soybean, reduced yield compared to ambient conditions even under elevated [CO₂]. This was largely due to heat stress on reproductive processes, especially during R5. Low‐intensity heat waves did not affect yields when applied during R1 but increased yields when applied during R5 likely due to relatively lower canopy temperatures and higher soil moisture, which uncoupled the negative effects of heating on cellular‐ and leaf‐level processes from plant‐level carbon assimilation. Modeling soybean yields based on carbon assimilation alone underestimated yield loss with high‐intensity heat waves and overestimated yield loss with low‐intensity heat waves, thus supporting the influence of direct heat stress on reproductive processes in determining yield. These results have implications for rain‐fed cropping systems and point toward a climatic tipping point for soybean yield when future heat waves exceed optimum temperature.     

Variation in vegetative and flowering phenology in a forest herb caused by environmental heterogeneity

Timing of seasonal plant development can affect biotic interactions and plant fitness. Phenology is governed largely by temperature and may therefore be affected by global climate warming, making this an important area of research. Several factors in addition to temperature may cause differences in phenology. We studied the influence of local environment, plant size, and reproductive effort on shoot emergence and flowering time of 290 individuals of Actaea spicata (Ranunculaceae), distributed among 25 plots in four populations. We used multiple regression and structural equation models (SEM) to study causal relationships. Among plots, soil temperature and canopy cover explained 63% of the variation in shoot emergence. Soil temperature, slope, and canopy cover together explained 83% of the variation in flowering time. Within plots, small plants on steep south-facing slopes with high soil potassium concentrations emerged earlier in the year. Plants emerging earlier flowered earlier, but no environmental factors affected flowering time directly. We found no effects of reproductive effort. Our results support the view that flowering time of temperate forest herbs is constrained by several environmental factors acting indirectly through effects on shoot emergence time.