High night temperature induces contrasting responses for spikelet fertility, spikelet tissue temperature, flowering characteristics and grain quality in rice

Climate change is increasing night temperature (NT) more than day temperature (DT) in rice-growing areas. Effects of combinations of NT (24-35°C) from microsporogenesis to anthesis at one or more DT (30 or 35°C) at anthesis on rice spikelet fertility, temperature within spikelets, flowering pattern, grain weight per panicle, amylose content and gel consistency were investigated in contrasting rice cultivars under controlled environments. Cultivars differed in spikelet fertility response to high NT, with higher fertility associated with cooler spikelets (P < 0.01). Flowering dynamics were altered by high NT and a novel high temperature tolerance complementary mechanism, shorter flower open duration in cv. N22, was identified. High NT reduced spikelet fertility, grain weight per panicle, amylose content and gel consistency, whereas high DT reduced only gel consistency. Night temperature >27°C was estimated to reduce grain weight. Generally, high NT was more damaging to grain weight and selected grain quality traits than high DT, with little or no interaction between them. The critical tolerance and escape traits identified, i.e. spikelet cooling, relatively high spikelet fertility, earlier start and peak time of anthesis and shorter spikelet anthesis duration can aid plant breeding programs targeting resilience in warmer climates.

Resilience of rice (Oryzaspp.) pollen germination and tube growth to temperature stress

Resilience of rice cropping systems to potential global climate change will partly depend on temperature tolerance of pollen germination (PG) and tube growth (PTG). Germination of pollen of high temperature susceptible Oryza glaberrima Steud. (cv. CG14) and O. sativa L. ssp. indica (cv. IR64) and high temperature tolerant O. sativa ssp. aus (cv. N22), was assessed on a 5.6-45.4°C temperature gradient system. Mean maximum PG was 85% at 27°C with 1488 μm PTG at 25°C. The hypothesis that in each pollen grain, minimum temperature requirements (Tn) and maximum temperature limits (Tx) for germination operate independently was accepted by comparing multiplicative and subtractive probability models. The maximum temperature limit for PG in 50% of grains (Tx(50)) was lowest (29.8°C) in IR64 compared with CG14 (34.3°C) and N22 (35.6°C). Standard deviation (sx) of Tx was also low in IR64 (2.3°C) suggesting that the mechanism of IR64's susceptibility to high temperatures may relate to PG. Optimum germination temperatures and thermal times for 1mm PTG were not linked to tolerating high temperatures at anthesis. However, the parameters Tx(50) and sx in the germination model define new pragmatic criteria for successful and resilient PG, preferable to the more traditional cardinal (maximum and minimum) temperatures.

Synergistic effects of geographical strain, temperature and larval food on insecticide tolerance in Callosobruchus maculatus (F.)

This study investigated the multifactorial interaction of various environmental factors including 17 geographical strain (Brazil, Cameroon and Yemen strains), temperature, dose and larval food 18 (cowpea and mungbean) on the response of Callosobruchus maculatus adult to insecticide. All 19 the main factors, their two-way interactions and the four-way interaction had significant effects 20 on C. maculatus response to malathion (an organophosphate insecticide). However, the three-21 way interactions were not statistically significant (except strain x food x dose, P = 0.002). The 22 2 Brazil strain was the most responsive to temperature irrespective of the larval food type. The 23 impact of food type differs from one strain to the other, for instance, the food that imparts higher 24 tolerance in a strain might reduce the tolerance in another. Likewise, the hierarchy of tolerance 25 among the cowpea-reared strains (Brazil > Cameroon > Yemen) was totally different from the 26 mungbean-reared strains (Cameroon > Yemen > Brazil). The reasons for these differences were 27 discussed in the light of their impact on C. maculatus management. The management of both C. 28 maculatus and development of resistance could be complex, hence, the states of a variety of 29 environmental factors need to be considered. This is necessary in order to maximize management 30 success of this bruchid especially in tropical/subtropical developing countries.

Effect of Rht alleles on the tolerance of wheat grain set to high temperature and drought stress during booting and anthesis

Factorial pot experiments were conducted to compare the responses of GA-sensitive and GA-insensitive reduced height (Rht) alleles in wheat for susceptibility to heat and drought stress during booting and anthesis. Grain set (grains/spikelet) of near isogenic lines (NILs) was assessed following three day transfers to controlled environments imposing day temperatures (t) from 20 to 40°C. Transfers were during booting and/or anthesis and pots maintained at field capacity (FC) or had water withheld. Logistic responses (y = c/1+e-b(t -m)) described declining grain set with increasing t, and t5 was that fitted to give a 5% reduction in grain set. Averaged over NIL, t5 for anthesis at FC was 31.7±0.47°C (S.E.M, 26 d.f.). Drought at anthesis reduced t5 by <2°C. Maintaining FC at booting conferred considerable resistance to high temperatures (t5=33.9°C) but booting was particularly heat susceptible without water (t5 =26.5°C). In one background (cv. Mercia), for NILs varying at the Rht-D1 locus, there was progressive reduction in t5 with dwarfing and reduced gibberellic acid (GA) sensitivity (Rht-D1a, tall, 32.7±0.72; Rht-D1b, semi-dwarf, 29.5±0.85; Rht-D1c, severe dwarf, 24.2±0.72). This trend was not evident for the Rht-B1 locus, or for Rht-D1b in an alternative background (Maris Widgeon). The GA-sensitive severe dwarf Rht12 was more heat tolerant (t5=29.4±0.72) than the similarly statured GA-insensitive Rht-D1c. The GA-sensitive, semi-dwarfing Rht8 conferred greater drought tolerance in one experiment. Despite the effects of Rht-D1 alleles in Mercia on stress tolerance, the inconsistency of the effects over background and locus led to the conclusion that semi-dwarfing with GA-insensitivity did not necessarily increase sensitivity to stress at booting and flowering. In comparison to effects of semi-dwarfing alleles, responses to heat stress are much more dramatically affected by water availability and the precise growth stage at which the stress is experienced by the plants.

Low-temperature-induced changes in trehalose, mannitol and arabitol associated with enhanced tolerance to freezing in ectomycorrhizal basidiomycetes ( Hebeloma spp.)

Ectomycorrhizal fungi have been shown to survive sub-zero temperatures in axenic culture and in the field. However, the physiological basis for resistance to freezing is poorly understood. In order to survive freezing, mycelia must synthesise compounds that pro-tect the cells from frost damage, and certain fungal-spe-cific soluble carbohydrates have been implicated in this role. Tissue concentrations of arabitol, mannitol and trehalose were measured in axenic cultures of eight Hebeloma strains of arctic and temperate origin grown at 22, 12, 6 and 2°C. In a separate experiment, mycelia were frozen to –5°C after pre-conditioning at either 2°C or 22°C. For some, especially temperate strains, there was a clear increase in specific soluble carbohydrates at lower growth temperatures. Trehalose and mannitol were present in all strains and the highest concentrations (close to 2.5% and 0.5% dry wt.) were recorded only after a cold period. Arabitol was found in four strains only when grown at low temperature. Cold pre-condi-tioning enhanced recovery of mycelia following freez-ing. In four out of eight strains, this was paralleled by increases in mannitol and trehalose concentration at low temperature that presumably contribute towards cryopro-tection. The results are discussed in an ecological con-text with regard to mycelial overwintering in soil.

Simulation of the impact of high temperature stress on annual crop yields

Brief periods of high temperature which occur near flowering can severely reduce the yield of annual crops such as wheat and groundnut. A parameterisation of this well-documented effect is presented for groundnut (i.e. peanut; Arachis hypogaeaL.). This parameterisation was combined with an existing crop model, allowing the impact of season-mean temperature, and of brief high-temperature episodes at various times near flowering, to be both independently and jointly examined. The extended crop model was tested with independent data from controlled environment experiments and field experiments. The impact of total crop duration was captured, with simulated duration being within 5% of observations for the range of season-mean temperatures used (20-28 degrees C). In simulations across nine differently timed high temperature events, eight of the absolute differences between observed and simulated yield were less than 10% of the control (no-stress) yield. The parameterisation of high temperature stress also allows the simulation of heat tolerance across different genotypes. Three parameter sets, representing tolerant, moderately sensitive and sensitive genotypes were developed and assessed. The new parameterisation can be used in climate change studies to estimate the impact of heat stress on yield. It can also be used to assess the potential for adaptation of cropping systems to increased temperature threshold exceedance via the choice of genotype characteristics. (c) 2005 Elsevier B.V. All rights reserved.

Heat tolerance in groundnut

Tolerance to high soil and air temperature during the reproductive phase is an important component of adaptation to and and semi-arid cropping environments in groundnut. Between 10 and 22 genotypes were screened for tolerance to high air and soil temperature in controlled environments. To assess tolerance to high soil temperature, 10 genotypes were grown from start of podding to harvest at ambient (28 degrees) and high (38 degreesC) soil temperatures, and crop growth rate (CGR), pod growth rate (PGR) and partitioning (ratio PGR:CGR) measured. To assess tolerance to high air temperature during two key stages-microsporogenesis (3-6 days before flowering, DBF) and flowering, fruit-set was measured in two experiments. In the first experiment, 12 genotypes were exposed to short (3-6 days) episodes of high (38 degreesC) day air temperature at 6 DBF and at flowering. In the second experiment, 22 genotypes were exposed to 40 degreesC day air temperature for I day at 6 DBF, 3 DBF or at flowering. Cellular membrane thermostability (relative injury, RI) was also measured in these 22 genotypes. There was considerable variation among genotypes in response to high temperature, whether assessed by growth rates, fruit-set or RI. Pod weight at high soil temperature was associated with variation in CGR rather than partitioning. Flowering was more sensitive to high air temperature than microsporogenesis. Genotypes tolerant to high air temperature at microsporogenesis were not necessarily tolerant at flowering, and nor was tolerance correlated with RI. Six genotypes (796, 55-437, ICG 1236, ICGV 86021, lCGV 87281 and ICGV 92121) were identified as heat tolerant based on their performance in all tests. These experiments have shown that groundnut genotypes can be easily screened for reproductive tolerance to high air and soil temperature and that several sources of heat tolerance are available in groundnut germplasm. (C) 2003 Elsevier Science B.V. All rights reserved.

High temperature stress and spikelet fertility in rice (Oryza sativa L.)

In future climates, greater heat tolerance at anthesis will be required in rice. The effect of high temperature at anthesis on spikelet fertility was studied on IR64 (lowland indica) and Azucena (upland Japonica) at 29.6 degrees C (control), 33.7 degrees C, and 36.2 degrees C tissue temperatures. The objectives of the study were to: (i) determine the effect of temperature on flowering pattern; (ii) examine the effect of time of day of spikelet anthesis relative to a high temperature episode on spikelet fertility; and (iii) study the interactions between duration of exposure and temperature on spikelet fertility. Plants were grown at 30/24 degrees C day/night temperature in a greenhouse and transferred to growth cabinets for the temperature treatments. Individual spikelets were marked with paint to relate fertility to the time of exposure to different temperatures and durations. In both genotypes the pattern of flowering was similar, and peak anthesis occurred between 10.30 h and 11.30 h at 29.2 degrees C, and about 45 min earlier at 36.2 degrees C. In IR64, high temperature increased the number of spikelets reaching anthesis, whereas in Azucena numbers were reduced. In both genotypes :511 h exposure to >= 33.7 degrees C at anthesis caused sterility. In IR64, there was no interaction between temperature and duration of exposure, and spikelet fertility was reduced by about 7% per degrees C > 29.6 degrees C. In Azucena there was a significant interaction and spikelet fertility was reduced by 2.4% degrees Cd-1 above a threshold of 33 degrees C. Marking individual spikelets is an effective method to phenotype genotypes and lines for heat tolerance that removes any apparent tolerance due to temporal escape.

Phenotyping parents of mapping populations of rice for heat tolerance during anthesis

Seed set of rice (Oryza sativa L.) is highly sensitive to short episodes of high temperature at anthesis events that are likely to be more frequent in future climates. Breeding for tolerance is therefore an essential component of adaptation to climate variability and change. Experiments were conducted in 2003 and 2004 at optimum (30 degrees C daytime) and high (35 and 38 degrees C) air temperature using parents of some prominent mapping populations (i) to determine whether there were differences in the daily flowering pattern and hence a potential heat avoidance mechanism, and (ii) to identify rice genotypes having true heat tolerance during anthesis, that is, high seed set in spikelets exposed to high temperature. Rice cultivar CG14 (O. glaberrima) reached peak anthesis earlier in the morning (1.5 h after dawn) under both control (30 degrees C) and high (38 degrees C) temperature conditions than O. sativa genotypes (>= 3 h after dawn). Exposure to high temperature (centered on the time of peak anthesis) for 6 h reduced spikelet fertility more than exposure for 2 h, and fertility was lower at 38 degrees C than at 35 degrees C. Genotypic ranking for spikelet fertility at 35 and 38 degrees C was highly correlated in both 2003 and 2004. Fertility was also highly correlated across years, suggesting a consistent and reproducible response of spikelet fertility to temperature. The check cultivar N22 was the most heat tolerant genotype (64-86% fertility at 38 degrees C) and cultivars Azucena and Moroberekan the most susceptible (<8%).

Analysis of shade and temperature effects on assimilate partitioning in the everbearing strawberry 'Everest' as the basis for optimised long-season fruit production

Growth patterns and cropping were evaluated over the season for the everbearing strawberry 'Everest' at a range of temperatures (15-27degreesC) in two light environments (ambient and 50% shade). The highest yield was recorded for unshaded plants grown at 23degreesC, but the optimum temperature for vegetative growth was 15degreesC. With increasing temperature fruit number increased, but fruit weight decreased. Fruit weight was also significantly reduced by shade, and although 'Everest' showed a degree of shade tolerance in vegetative growth, yield was consistently reduced by shade. Shade also reduced the number of crowns developed by the plants over the course of the season, emphasising that crown number was ultimately the limiting factor for yield potential. We conclude that, in contrast to Junebearers which partition more assimilates to fruit at temperatures around 15degreesC (Le Miere et al., 1998), optimised cropping in the everbearer 'Everest' is achieved at the significantly higher temperature of 23degreesC. These findings have significance for commercial production, in which protection tends to reduce light levels but increase average temperature throughout the season.

Physiological and proteomic approaches to address heat tolerance during anthesis in rice (Oryza sativa L.)

Episodes of high temperature at anthesis, which in rice is the most sensitive stage to temperature, are expected to occur more frequently in future climates. The morphology of the reproductive organs and pollen number, and changes in anther protein expression, were studied in response to high temperature at anthesis in three rice (Oryza sativa L.) genotypes. Plants were exposed to 6 h of high (38 °C) and control (29 °C) temperature at anthesis and spikelets collected for morphological and proteomic analysis. Moroberekan was the most heat-sensitive genotype (18% spikelet fertility at 38 °C), while IR64 (48%) and N22 (71%) were moderately and highly heat tolerant, respectively. There were significant differences among the genotypes in anther length and width, apical and basal pore lengths, apical pore area, and stigma and pistil length. Temperature also affected some of these traits, increasing anther pore size and reducing stigma length. Nonetheless, variation in the number of pollen on the stigma could not be related to measured morphological traits. Variation in spikelet fertility was highly correlated (r=0.97, n=6) with the proportion of spikelets with ≥20 germinated pollen grains on the stigma. A 2D-gel electrophoresis showed 46 protein spots changing in abundance, of which 13 differentially expressed protein spots were analysed by MS/MALDI-TOF. A cold and a heat shock protein were found significantly up-regulated in N22, and this may have contributed to the greater heat tolerance of N22. The role of differentially expressed proteins and morphology during anther dehiscence and pollination in shaping heat tolerance and susceptibility is discussed.

Genetic analysis of heat tolerance at anthesis in rice

Genetic analysis of heat tolerance will help breeders produce rice (Oryza sativa L.) varieties adapted to future climates. An F6 population of 181 recombinant inbred lines of Bala (tolerant) × Azucena (susceptible) was screened for heat tolerance at anthesis by measuring spikelet fertility at 30°C (control) and 38°C (high temperature) in experiments conducted in the Philippines and the United Kingdom. The parents varied significantly for absolute spikelet fertility under control (79–87%) and at high temperature (2.9–47.1%), and for relative spikelet fertility (high temperature/control) at high temperature (3.7–54.9%). There was no correlation between spikelet fertility in control and high-temperature conditions and no common quantitative trait loci (QTLs) were identified. Two QTLs for spikelet fertility under control conditions were identified on chromosomes 2 and 4. Eight QTLs for spikelet fertility under high-temperature conditions were identified on chromosomes 1, 2, 3, 8, 10, and 11. The most significant heat-responsive QTL, contributed by Bala and explaining up to 18% of the phenotypic variation, was identified on chromosome 1 (38.35 mega base pairs on the rice physical genome map). This QTL was also found to influence plant height, explaining 36.6% of the phenotypic variation. A comparison with other studies of abiotic (drought, cold, salinity) stresses showed QTLs at similar positions on chromosomes 1, 3, 8, and 10, suggesting common underlying stress-responsive regions of the genome.

Rice seed quality development and temperature during late development and maturation

The potential longevity of japonica rice (Oryza sativa L. subsp. japonica) seed is particularly sensitive to high temperature – and thus climate change – during development and maturation. Cultivar Taipei 309 was grown at 28/208C (12 h/12 h) and then from 19 DAA (days after 50% anthesis), when seeds were just over half filled, at 28/208C, 30/228C, 32/248C or 34/268C (12 h/12 h). Whereas ability to germinate ex planta had been achieved in almost all seeds by 24 DAA, only half the population were desiccation tolerant. Desiccation tolerance continued to increase over the subsequent 28 d, similarly at all four temperatures. Subsequent longevity, assessed by p50 (period in days to reduce viability to 50% in hermetic storage at 408C with c. 15% moisture content), increased progressively at 28/208C until 38 DAA, and remained constant until the final harvest (52 DAA). The three warmer temperature regimes provided similar longevity to 28/208C at any one harvest, except at 38 DAA where the warmest (34/268C) was poorer. That temperature regime also provided greater seed-to-seed variability within each survival curve. The results confirm that appreciable improvement in seed quality occurs during seed development and also subsequent maturation in japonica rice, but that increase in temperature from 28/208C to 34/268C during late seed filling onwards has comparatively little effect thereon. Comparison with previous investigations suggests that seed quality development may be less sensitive to high temperatures during late development and maturation than during the early seed development that precedes it.

Legume type and temperature effects on the toxicity of insecticide to the genus Callosobruchus (Coleoptera: Bruchidae)

Three bruchid pest species, Callosobruchus maculatus, Callosobruchus chinensis and Callosobruchus rhodesianus, were studied for their response to insecticide toxicity taking into account the separate and interactive effects of temperature and pre-adult food. The food types used were cowpea (Vigna unguiculata) and mungbean (Vigna radiata). Callosobruchus maculatus was the most tolerant to malathion and the least affected by temperature change while C. rhodesianus was the least tolerant. Over a 4 C range (23, 25, 27 C), there was generally a significant impact of temperature on the tolerance of the three species to the insecticide. The food type on which the insects developed influenced considerably the degree of insecticide tolerance. Callosobruchus maculatus and C. chinensis populations reared onmungbean had higher tolerance to malathion than their counterparts reared on cowpea, but the opposite was observed in C. rhodesianus populations. The food influence in this study suggested an ancestral cause or fitness cost depending on the species. The interaction of food-by-temperature had no significant effect on malathion toxicity to this genus. Correlation analysis showed C. chinensis to be relatively less sensitive to insecticide concentration over the range studied compared with the other two species.

Varietal effects of cowpea, Vigna unguiculata, on tolerance to malathion in Callosobruchus maculatus (Coleoptera: Bruchidae)

The impact of cowpea variety on the response of cowpea bruchid, Callosobruchus maculatus, to malathion was investigated. The interaction of six cowpea varieties (Adamawa Brown, Ife BPC, Ife Brown, Lilongwe, Ntcheu and NCRI-L25) with the geographical strains of C. maculatus (Brazil and Cameroon), temperature (23, 25, 27 C) and insecticide concentration were considered. Cowpea variety (V) had an unpredictable effect on C. maculatus response to malathion. Bruchid populations produced by Ife BPC were the most susceptible to malathion while those yielded by NCRI-L25 were the most tolerant. Regardless of the cowpea variety, the Brazil strain showed higher tolerance than the Cameroon strain. There was significant effect of temperature (T) and insecticide concentration (C) on malathion tolerance in both strains (S). Likewise, there was significant impact of all two-way interactions on cowpea bruchid tolerance except V x C. Significant three-way interactions on C. maculatus tolerance to malathion was only observed in S T V and S T C. The predictability of changing one of the factors on the susceptibility of C. maculatus to insecticide was very low. This study suggests a need to take the insecticide tolerance of insect populations produced by novel varieties into account during plant breeding in addition to factors such as yield and resistance to insect and disease attack.