RESUMO
The escalation in the intensity, frequency, and duration of high-temperature (HT) stress is currently unparalleled, which aggravates the challenges for crop production. Yet, the stage-dependent responses of reproductive organs to HT stress at the morphological, physiological, and molecular levels remain inadequately explored in pivotal staple crops. This review synthesized current knowledge regarding the mechanisms by which HT stress induces abnormalities and aberrations in reproductive growth and development, as well as by which it alters the morphology and function of florets, flowering patterns, and the processes of pollination and fertilization in maize (Zea mays L.). We identified the stage-specific sensitivities to HT stress and accurately defined the sensitive period from a time scale of days to hours. The microspore tetrad phase of pollen development and anthesis (especially shortly after pollination) are most sensitive to HT stress, and even brief temperature spikes during these stages can lead to significant kernel loss. The impetuses behind the heat-induced impairments in seed set are closely related to carbon, reactive oxygen species, phytohormone signals, ion (e.g. Ca2+) homeostasis, plasma membrane structure and function, and others. Recent advances in understanding the genetic mechanisms underlying HT stress responses during maize sexual reproduction have been systematically summarized.
Assuntos
Resposta ao Choque Térmico , Reprodução , Zea mays , Zea mays/fisiologia , Zea mays/crescimento & desenvolvimento , Resposta ao Choque Térmico/fisiologia , Temperatura Alta , Pólen/fisiologiaRESUMO
In a warming climate, high temperature stress greatly threatens crop yields. Maize is critical to food security, but frequent extreme heat events coincide temporally and spatially with the period of kernel number determination (e.g., flowering stage), greatly limiting maize yields. In this context, how to increase or at least maintain maize yield has become more important. Nitrogen fertilizer (N) is widely used to improve maize yields, but its effect in heat stress is unclear. For this, we collected 1536 pairs of comparisons from 113 studies concerning N conducted in the past 20 years over China. We classified the data into two groups - without high temperature stress (NHT) and with high temperature stress during the critical period for maize kernel number determination (HT) - based on the national meteorological data. We comprehensively evaluated N effects on grain yield under HT and NHT using meta-analysis. The effect of N on maize yield became significantly smaller in HT than that in NHT. In NHT, soil characteristics, crop management practices, and climatic conditions all significantly affected N effects on maize yield, but in HT, only a few factors such as soil organic matter and mean annual precipitation significantly affected N effects. Hence, it is difficult to improve N effect by improving soil characteristics and crop management when meeting with high temperature stress during flowering. On average, N effect increased with increased N input, but there were respective N input thresholds in NHT and HT, beyond which N effects on maize yield remained stable. According to the thresholds, it is speculated that moderately reducing N input (~20 %) likely increased high temperature tolerance of maize during flowering. These findings have important implications for the optimization of N management under a warming climate.
Assuntos
Nitrogênio , Zea mays , Zea mays/fisiologia , Zea mays/crescimento & desenvolvimento , China , Fertilizantes , Temperatura Alta , Resposta ao Choque Térmico/fisiologia , Mudança ClimáticaRESUMO
Heat waves induced by climate warming have become common in food-producing regions worldwide, frequently coinciding with high temperature (HT)-sensitive stages of many crops and thus threatening global food security. Understanding the HT sensitivity of reproductive organs is currently of great interest for increasing seed set. The responses of seed set to HT involve multiple processes in both male and female reproductive organs, but we currently lack an integrated and systematic summary of these responses for the world's three leading food crops (rice, wheat, and maize). In the present work, we define the critical high temperature thresholds for seed set in rice (37.2°C ± 0.2°C), wheat (27.3°C ± 0.5°C), and maize (37.9°C ± 0.4°C) during flowering. We assess the HT sensitivity of these three cereals from the microspore stage to the lag period, including effects of HT on flowering dynamics, floret growth and development, pollination, and fertilization. Our review synthesizes existing knowledge about the effects of HT stress on spikelet opening, anther dehiscence, pollen shedding number, pollen viability, pistil and stigma function, pollen germination on the stigma, and pollen tube elongation. HT-induced spikelet closure and arrest of pollen tube elongation have a catastrophic effect on pollination and fertilization in maize. Rice benefits from pollination under HT stress owing to bottom anther dehiscence and cleistogamy. Cleistogamy and secondary spikelet opening increase the probability of pollination success in wheat under HT stress. However, cereal crops themselves also have protective measures under HT stress. Lower canopy/tissue temperatures compared with air temperatures indicate that cereal crops, especially rice, can partly protect themselves from heat damage. In maize, husk leaves reduce inner ear temperature by about 5°C compared with outer ear temperature, thereby protecting the later phases of pollen tube growth and fertilization processes. These findings have important implications for accurate modeling, optimized crop management, and breeding of new varieties to cope with HT stress in the most important staple crops.