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.

Effect of elevated CO2 and high temperature on seed-set and grain quality of rice

Hybrid vigour may help overcome the negative effects of climate change in rice. A popular rice hybrid (IR75217H), a heat-tolerant check (N22), and a mega-variety (IR64) were tested for tolerance of seed-set and grain quality to high-temperature stress at anthesis at ambient and elevated [CO2]. Under an ambient air temperature of 29 °C (tissue temperature 28.3 °C), elevated [CO2] increased vegetative and reproductive growth, including seed yield in all three genotypes. Seed-set was reduced by high temperature in all three genotypes, with the hybrid and IR64 equally affected and twice as sensitive as the tolerant cultivar N22. No interaction occurred between temperature and [CO2] for seed-set. The hybrid had significantly more anthesed spikelets at all temperatures than IR64 and at 29 °C this resulted in a large yield advantage. At 35 °C (tissue temperature 32.9 °C) the hybrid had a higher seed yield than IR64 due to the higher spikelet number, but at 38 °C (tissue temperature 34–35 °C) there was no yield advantage. Grain gel consistency in the hybrid and IR64 was reduced by high temperatures only at elevated [CO2], while the percentage of broken grains increased from 10% at 29 °C to 35% at 38 °C in the hybrid. It is concluded that seed-set of hybrids is susceptible to short episodes of high temperature during anthesis, but that at intermediate tissue temperatures of 32.9 °C higher spikelet number (yield potential) of the hybrid can compensate to some extent. If the heat tolerance from N22 or other tolerant donors could be transferred into hybrids, yield could be maintained under the higher temperatures predicted with climate change.

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.

MELHORAMENTO do TRIGO: XXXI. HERDABILIDADES E CORRELAÇÕES ENTRE TRÊS CARACTERÍSTICAS AGRONÔMICAS em POPULAÇÕES HÍBRIDAS

Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)

Yield of upland rice cultivars in rainfed and sprinkler-irrigated systems in the Cerrado region of Brazil

In upland rice, the reduced grain yield that originates from the moisture stress period is related to the plant's tolerance of water deficiency and may vary between cultivars. The purpose of the work presented here was to evaluate the performance of upland rice cultivars in both rainfed and sprinkler-irrigated systems. A split-plot scheme with 8 replicates in a randomised block design was used. The plots were composed of 2 cropping systems (rainfed and sprinkler-irrigated) and the split-plot consisted of 2 cultivars (IAC 201 and Carajas) which are suggested for cultivation in upland ecosystems. Carajas had a greater number of panicles per square metre, higher spikelet fertility and grain mass, and, consequently, a higher grain yield than IAC 201 regardless of cropping system. IAC 201 was more sensitive to water deficiency than Carajas. Sprinkler irrigation improved spikelet fertility, grain mass and upland rice grain yield. Even when cultivars with a higher tolerance of water deficiency are used, a sprinkler-irrigated system may be a viable method to increase upland rice yield throughout most of the Brazilian Cerrado.

Nitrogen and silicon fertilization of upland rice

O silício não é considerado um elemento essencial para o crescimento e desenvolvimento das plantas, entretanto, sua absorção traz inúmeros benefícios, principalmente ao arroz, como aumento da espessura da parede celular, conferindo resistência mecânica a penetração de fungos, melhora o ângulo de abertura das folhas tornando-as mais eretas, diminuindo o auto-sombreamento e aumentando a resistência ao acamamento, especialmente sob altas doses de nitrogênio. O presente trabalho teve por objetivo avaliar os efeitos da adubação nitrogenada e silicatada nos componentes vegetativos, nos componentes da produção, na altura da planta e na produtividade da cultivar de arroz IAC 202. O experimento foi constituído da combinação de três doses de nitrogênio (5, 75 e 150 mg de N kg-1 de solo) aplicado na forma de uréia e quatro doses de silício (0, 200, 400 e 600 mg de SiO2 kg-1 de solo), aplicado na forma de silicato de cálcio. O delineamento experimental utilizado foi o inteiramente casualizado em esquema fatorial 3 ´ 4 (N = 5). A adubação nitrogenada aumentou o número de colmos e panículas por metro quadrado e o número total de espiguetas, refletindo na produtividade de grãos. O perfilhamento excessivo causado pela adubação nitrogenada inadequada causou redução na porcentagem de colmos férteis, na fertilidade das espiguetas e da massa de grãos. A adubação silicatada reduziu o número de espiguetas chochas por panícula e aumentou a massa de grãos sem, contudo, refletir na produtividade de grãos.

Fertilizer distribution mechanisms and side dress nitrogen fertilization in upland rice under no-tillage system

Algumas culturas têm pouca adaptação ao sistema plantio direto (SPD), em vista da alta compactação da camada superficial do solo. Nesse caso, o mecanismo utilizado na semeadora para a abertura dos sulcos para deposição do fertilizante pode ter grande importância no sentido de facilitar a penetração das raízes. Avaliouse a influência do mecanismo de distribuição de fertilizante e da adubação nitrogenada na cultura do arroz de terras altas (Oryza sativa) no sistema plantio direto. O experimento foi conduzido nos anos agrícolas 2001/2002 e 2002/2003, em Botucatu-SP, Brasil. O delineamento experimental foi em blocos casualizados, com parcelas subdivididas e quatro repetições. As parcelas foram constituídas por dois mecanismos de distribuição de fertilizantes (haste sulcadora e disco duplo). Nas subparcelas, quatro níveis de N foram aplicados em cobertura (0, 40, 80 e 120 kg ha-1). Avaliou-se a profundidade de abertura do sulco e de deposição das sementes, a população de plantas, a altura de plantas, o número de colmos e de panículas m-2, o número total de espiguetas por panícula, a fertilidade das espiguetas, a massa de 1.000 grãos, a matéria seca da parte aérea, a produtividade e o teor de N na folha bandeira. O sucesso no estabelecimento da cultura do arroz de terras altas no SPD nas regiões de inverno seco do Brasil está diretamente ligado ao mecanismo de distribuição das semeadoras-adubadoras. A haste sulcadora promoveu maior profundidade de deposição de sementes, conseqüentemente reduzindo o estande, o número de panículas por área e a produtividade. A aplicação de N em cobertura no arroz de terras altas em SPD proporciona maiores produtividades quando a semeadura é realizada com mecanismo de disco duplo.

Manejo de irrigação por aspersão com base no Kc e adubação mineral na cultura de arroz de terras altas

A irrigação por aspersão diminui bastante o risco de perda da lavoura por deficiência hídrica e aumenta a produtividade de grãos, incentivando maior uso de tecnologias como adubação mineral. Com o objetivo de avaliar o efeito de diferentes manejos da água da irrigação por aspersão com base no coeficiente de cultura (Kc) e da adubação mineral sobre a cultura do arroz cv. IAC 201, foram instalados dois experimentos em Latossolo Vermelho Distrófico, em Selvíria (MS). O delineamento foi em blocos casualizados, com quatro repetições. Os tratamentos constituíram-se de precipitação pluvial natural e três manejos de água fornecidos por aspersão. O manejo (M2) foi realizado com base no Kc do arroz de terras altas. Os manejos M1 e M3 foram definidos como 0,5 e 1,5 vezes os Kcs utilizados em M2 respectivamente. em 1995/96, utilizou-se o esquema de parcelas subdivididas, sendo as subparcelas constituídas por dois níveis de adubação: AD1 - 12 kg de N, 90 kg de P2O5 e 30 kg de K2O ha-1, e AD2 - 24 kg de N, 180 kg de P2O5 e 60 kg de K2O ha-1. A deficiência hídrica da emergência da plântula até a diferenciação do primórdio da panícula provocou aumento do ciclo e redução do porte da planta. A deficiência hídrica entre os estádios de diferenciação do primórdio da panícula e os de emborrachamento reduziu o número de espiguetas por panícula. A utilização de 1,5 vezes os valores de Kc recomendados, no manejo da irrigação por aspersão proporcionou maior produtividade de grãos. Os níveis de adubação utilizados não influenciaram a resposta da cultura ao manejo da irrigação por aspersão.

Resposta de cultivares de arroz a doses de nitrogênio e do regulador de crescimento cloreto de clormequat

O objetivo deste trabalho foi avaliar os efeitos de doses de nitrogênio (0, 50, 100 e 150 kg ha-1) e do regulador de crescimento cloreto de clormequat (0, 1 e 2 L ha-1), em algumas características agronômicas de cultivares de arroz IAC 201 e IAC 202. O delineamento experimental foi o de blocos ao acaso, com quatro repetições. O N foi aplicado em cobertura, no estádio da diferenciação floral, e o regulador de crescimento foi aplicado parceladamente, aos 20 e 30 dias após a emergência das plântulas de arroz (perfilhamento). Na safra 2001/2002, houve efeito significativo de N para altura de plantas, comprimento da panícula, espiguetas por panícula, massa de 100 grãos e produtividade de grãos. Houve efeito do regulador de crescimento sobre o comprimento da panícula e espiguetas por panícula. A maior produtividade foi a da cultivar IAC 202. Na safra 2002/2003, houve efeito de N para altura de plantas, massa de 100 grãos, fertilidade das espiguetas e produtividade de grãos. O regulador de crescimento não exerceu efeito nas características testadas, e a cultivar IAC 202 foi novamente a mais produtiva.

Aplicação foliar e em cobertura de nitrogênio na cultura do trigo no cerrado

The nitrogen in wheat is essential for obtaining high yields, not only the dose but also the time and the way of application are critical, reducing potential leaching and the cost of production. The objective is evaluating leaf and sidedressing nitrogen application on wheat crop in years of 2006 and 2007. A randomized blocks design in a factorial scheme 5x3x2 was used. The treatments consisted of five doses of nitrogen in the solution (0; 2.5; 5.0; 7.5 and 10%), three application times (at tillering: 30 days after plant emergency (DAE), at full flowering (50 DAE) + in the beginning of grain formation (70 DAE) and at tillering + in the beginning of grain formation), with and without sidedressing nitrogen applied at 40 DAE, using urea as source. They were evaluated: chlorophyll and nitrogen content in leaf, number of spikelets per ear, number of grains per ear, mass of grains per ear, number of grains per spikelet, mass hectolitric, mass of 100 grains and productivity of grains. The application of nitrogen topdressing in both years, influenced the yield characteristics of wheat. The times of leaf nitrogen only affected the leaf N content. The leaf nitrogen concentrations increased linearly the number of grains per spikelets, grains per spike, chlorophyll content, grain weight per ear and grain yield, and reduced mean weight per hectoliter, only in 2007.

Maturação de sementes de aveia-preta (Avena strigosa Schreb). II. Maturidade da panícula e da espigueta

The present work aims at defining the panicle and spikelet maturity of Avena strigosa Schreb. Four experiments were carried out under field conditions during 1989 (E1 and E2) and 1990 (E3 and E4) in Botucatu county, São Paulo State, Brazil. The sowings were made in 04.19.89 (E1) and 06.18.89 (E2). Maturity of panicle was evaluated by weekly (E1) or biweekly harvests (E2) beginning at total emergence of the panicle. In E3 and E4, sowing was made in 06.04.90 and each experiment had a single harvest date (E3 = 11.01.90 and E4 = 11.07.90). After harvest, the spikelets with similar stage of development were grouped and evaluated. Based on dry matter accumulation and germination percentage of the seeds, it was observed that the maturity of the seeds in the panicle was reached at 28 and 35 days after total emergence of the panicle in the April sowing, and between 24.5 and 28 days in the June sowing. During this period a few percentage of panicles not yet yellow became all yellow, the cariopsis at semi-hard stage became hard and moisture of the seeds dropped from 32% to 12%. The maturity of the seeds in the spikelet was reached when the glumes changed their color from yellowish to yellow, the lemma and palea changed from yellow with black strips to black, the cariopsis changed from farinaceous stage to semi-hard and the moisture content of the seeds changed from 40% to 25%.

Floral development and embryology in the early-divergent grass Pharus

A detailed study of floral ontogeny, anatomy, and embryology in two (of six) species of Pharus is presented as part of a series of comparative investigations on early-divergent grasses. Pharus is a taxonomically isolated genus belonging to the earliest-diverging grass lineage with a true grass spikelet. It is unusual in possessing remarkably dimorphic florets: male florets possess two lodicules, six stamens, and a pistillode, whereas female florets lack lodicules entirely but possess six staminodes and a tricarpellary ovary with three stigmas. The rudimentary lodicules in male florets are initiated after the stamen whorls. There are most commonly six androecial organs, but in some florets, a five-staminate condition was observed, resulting from suppression of the abaxial stamen from the inner whorl, or even a four-staminate condition resulting from subsequent fusion of the two adaxial outer stamens (i.e., elements of both whorls). Thus, the pattern of floral zygomorphy in Pharus differs from that of many other grasses. Centrifixed anther attachment is reported for the first time in Pharus, resembling the condition in another early-divergent grass, Anomochloa, though anthers are introrse in Anomochloa compared with latrorse in Pharus. Anther wall development is of the reduced type in Pharus, in contrast to most other monocots. Microsporogenesis is of the successive type, as in many other monocots. The ovary develops from three distinct primordia and is unilocular with a single ovule and a pronounced ovary beak that is highly characteristic of Pharus. There is a hollow style, in contrast to the solid styles that are common in many other grasses. The embryo is highly differentiated, as in other grasses, with a distinct epiblast and a small cleft between the scutellum and the coleorhiza.

Maturação de sementes de aveia-preta (Avena strigosa Schreb). I. Maturidade do campo

The present work aimed to define the field maturity of Avena strigosa Schreb seeds cv. Comum. Three experiments were carried out in field conditions during 1986, 1987 and 1989 in Botucatu, SP, Brazil. The sowings were made in May. The characteristics evaluated were: panicles and spikelet aspects, the weight of the dry matter of 100 seeds with and without glumes, the seed moisture, the percentage of germinated seeds (sum of normal + abnormal seedlings), the percentage of germination and vigor (first count of germination test). Based on the dry matter accumulation and the gemination percentage of the seeds, it was concluded that the field maturity of the seeds was reached around 28 and 3 5 days after 100% of panicles emergence of the field. At this stage, the field presented a few percentage of particles (+/- 20%) not yet all yellowish, with spikelets at semi-hard stage and the seeds with 20 to 30% of moisture.

Homologies of the flower and inflorescence in the early-divergent grass anomochloa (poaceae)

Premise of the study: The grass subfamily Anomochlooideae is phylogenetically significant as the sister group to all other grasses. Thus, comparison of their structure with that of other grasses could provide clues to the evolutionary origin of these characters. Methods: We describe the structure, embryology, and development of the flower and partial inflorescence of the monotypic Brazilian grass Anomochloa marantoidea. We compare these features with those of other early-divergent grasses such as Pharus and Streptochaeta and closely related Poales such as Ecdeiocolea. Key results: Anomochloa possesses several features that are characteristic of Poaceae, notably a scutellum, a solid style, reduced stamen number, and an ovary with a single ovule that develops into a single indehiscent fruit. Interpretation of floral patterning in Anomochloa is problematic because the ramification pattern of the florets places the bracts and axes in unusual positions relative to the primary inflorescence axis. Our study indicates that there is a single abaxial carpel in Anomochloa, probably due to a cryptic type of pseudomonomery in Anomochloa that resembles the pseudomonomery of other grasses. On the other hand, the Anomochloa flower differs from the typical grass flower in lacking lodicules and possessing four stamens, in contrast with the tristaminate condition that characterizes many other grasses. Conclusions: Using the median part of the innermost bract as a locator, we tentatively homologize the inner bract of the Anomochloa partial inflorescence with the palea of other grasses. In this interpretation, the pattern of monosymmetry due to stamen suppression differs from that of Ecdeiocolea. © 2012 Botanical Society of America.