Evaluating the underlying physiological and molecular mechanisms in the system of rice intensification performance with Trichoderma-rice plant symbiosis as a model system.

The system of rice intensification (SRI) is an extensively-researched and increasingly widely-utilized methodology for alleviating current constraints on rice production. Many studies have shown physiological and morphological improvements in rice plants induced by SRI management practices to be very similar to those that are associated with the presence of beneficial microbial endophytes in or around rice plants, especially their roots. With SRI methods, grain yields are increased by 25-100% compared to conventional methods, and the resulting plant phenotypes are better able to cope with biotic and abiotic stresses. SRI management practices have been shown to be associated with significant increases in the populations of certain microorganisms known to enhance soil health and plant growth, e.g., Azospirillum, Trichoderma, Glomus, and Pseudomonas. This article evaluates the effects of applying Trichoderma as a model microbe for assessing microbial growth-promotion, biological control activity, and modulation of gene expression under the conditions created by SRI practices. Information about the molecular changes and interactions associated with certain effects of SRI management suggests that these practices are enhancing rice plants' expression of their genetic potentials. More systematic studies that assess the effects of SRI methods respectively and collectively, compared with standard rice production methods, are needed to develop a more encompassing understanding of how SRI modifications of crops' growing environment elicit and contribute to more robust and more productive phenotypes of rice.

Microbial Contributions for Rice Production: From Conventional Crop Management to the Use of 'Omics' Technologies.

Rice, the main staple food for about half of the world's population, has had the growth of its production stagnate in the last two decades. One of the ways to further improve rice production is to enhance the associations between rice plants and the microbiome that exists around, on, and inside the plant. This article reviews recent developments in understanding how microorganisms exert positive influences on plant growth, production, and health, focusing particularly on rice. A variety of microbial species and taxa reside in the rhizosphere and the phyllosphere of plants and also have multiple roles as symbiotic endophytes while living within plant tissues and even cells. They alter the morphology of host plants, enhance their growth, health, and yield, and reduce their vulnerability to biotic and abiotic stresses. The findings of both agronomic and molecular analysis show ways in which microorganisms regulate the growth, physiological traits, and molecular signaling within rice plants. However, many significant scientific questions remain to be resolved. Advancements in high-throughput multi-omics technologies can be used to elucidate mechanisms involved in microbial-rice plant associations. Prospectively, the use of microbial inoculants and associated approaches offers some new, cost-effective, and more eco-friendly practices for increasing rice production.

Topsoil microbial community structure responds to land cover type and environmental zone in the Western Pacific region.

Soil encompasses diverse microbial communities that are essential for fundamental ecosystem functions such as biogeochemical cycling. To better understand underlying biogeochemical processes, it is necessary to know the structure of soil archaeal and bacterial communities and their responses to edaphic and climate variables within and across various land cover types (LCTs) and environmental zones (ENZs). Here we sampled eighty-nine sites across five ENZs and four LCTs within the Western Pacific region. Through leveraging the second-generation sequencing of topsoil samples, we showed that α-diversity (taxonomic diversity) of archaea strongly varied within LCTs, whereas bacterial α-diversity was significantly controlled by both LCT and ENZ. Soil archaea and bacteria showed global niche differentiation associated with contrasting diversity responses to latitude and differential responses of microbial diversity patterns to edaphic and climate variables within LCTs and ENZs. In contrast to α-diversity, microbial ß-diversity (the compositional dissimilarity between sites) was majorly governed by ENZs, particularly for archaea (P < 0.01). Our results highlight the importance of LCTs and ENZs for understanding soil microbial contributions to nutrient dynamics and ecosystem resilience under land-use intensification and climate change.

Benefits to Plant Health and Productivity From Enhancing Plant Microbial Symbionts.

Plants exist in close association with uncountable numbers of microorganisms around, on, and within them. Some of these endophytically colonize plant roots. The colonization of roots by certain symbiotic strains of plant-associated bacteria and fungi results in these plants performing better than plants whose roots are colonized by only the wild populations of microbes. We consider here crop plants whose roots are inhabited by introduced organisms, referring to them as Enhanced Plant Holobionts (EPHs). EPHs frequently exhibit resistance to specific plant diseases and pests (biotic stresses); resistance to abiotic stresses such as drought, cold, salinity, and flooding; enhanced nutrient acquisition and nutrient use efficiency; increased photosynthetic capability; and enhanced ability to maintain efficient internal cellular functioning. The microbes described here generate effects in part through their production of Symbiont-Associated Molecular Patterns (SAMPs) that interact with receptors in plant cell membranes. Such interaction results in the transduction of systemic signals that cause plant-wide changes in the plants' gene expression and physiology. EPH effects arise not only from plant-microbe interactions, but also from microbe-microbe interactions like competition, mycoparasitism, and antibiotic production. When root and shoot growth are enhanced as a consequence of these root endophytes, this increases the yield from EPH plants. An additional benefit from growing larger root systems and having greater photosynthetic capability is greater sequestration of atmospheric CO2. This is transferred to roots where sequestered C, through exudation or root decomposition, becomes part of the total soil carbon, which reduces global warming potential in the atmosphere. Forming EPHs requires selection and introduction of appropriate strains of microorganisms, with EPH performance affected also by the delivery and management practices.

Symbiotic Root-Endophytic Soil Microbes Improve Crop Productivity and Provide Environmental Benefits.

Plants should not be regarded as entities unto themselves, but as the visible part of plant-microbe complexes which are best understood as "holobiomes." Some microorganisms when given the opportunity to inhabit plant roots become root symbionts. Such root colonization by symbiotic microbes can raise crop yields by promoting the growth of both shoots and roots, by enhancing uptake, fixation, and/or more efficient use of nutrients, by improving plants' resistance to pests, diseases, and abiotic stresses that include drought, salt, and other environmental conditions, and by enhancing plants' capacity for photosynthesis. We refer plant-microbe associations with these capabilities that have been purposefully established as enhanced plant holobiomes (EPHs). Here, we consider four groups of phylogenetically distinct and distant symbiotic endophytes: (1) Rhizobiaceae bacteria; (2) plant-obligate arbuscular mycorrhizal fungi (AMF); (3) selected endophytic strains of fungi in the genus Trichoderma; and (4) fungi in the Sebicales order, specifically Piriformospora indica. Although these exhibit quite different "lifestyles" when inhabiting plants, all induce beneficial systemic changes in plants' gene expression that are surprisingly similar. For example, all induce gene expression that produces proteins which detoxify reactive oxygen species (ROS). ROS are increased by environmental stresses on plants or by overexcitation of photosynthetic pigments. Gene overexpression results in a cellular environment where ROS levels are controlled and made more compatible with plants' metabolic processes. EPHs also frequently exhibit increased rates of photosynthesis that contribute to greater plant growth and other capabilities. Soil organic matter (SOM) is augmented when plant root growth is increased and roots remain in the soil. The combination of enhanced photosynthesis, increasing sequestration of CO2 from the air, and elevation of SOM removes C from the atmosphere and stores it in the soil. Reductions in global greenhouse gas levels can be accelerated by incentives for carbon farming and carbon cap-and-trade programs that reward such climate-friendly agriculture. The development and spread of EPHs as part of such initiatives has potential both to enhance farm productivity and incomes and to decelerate global warming.

Roles of microbes in supporting sustainable rice production using the system of rice intensification.

The system of rice intensification (SRI) is an agroecological approach to rice cultivation that seeks to create optimal conditions for healthy plant growth by minimizing inter-plant competition, transplanting widely spaced young single seedlings, and optimizing favorable soil conditions with organic amendments, increased soil aeration by weeding, and controlled water management. These practices improve rice plant growth with yields up to three times more than with conventional cultivation methods, and increase crop resilience under biotic and abiotic stresses. This review discusses the roles of beneficial microbes in improving rice plant growth, yield, and resilience when SRI practices are used, and how these modifications in plant, soil, water, and nutrient management affect the populations and diversity of soil microorganisms. Mechanisms whereby symbiotic microbes support rice plants' growth and performance are also discussed.

Effects of Trichoderma seedling treatment with System of Rice Intensification management and with conventional management of transplanted rice.

Many benefits of Trichoderma inoculation for improving crop production have been documented, including growth and yield enhancement and the alleviation of biotic and abiotic stresses. However, because rice is usually cultivated under continuous flooding that creates anaerobic soil conditions, this limits the benefits of these beneficial fungi. Cultivating rice with the methods of the System of Rice Intensification (SRI) provides rice plants with a more favorable environment for their colonization by beneficial microbes in the soil because the soil is more aerobic under SRI management and contains more organic matter. This study evaluated the effects of Trichoderma inoculation of rice plants under SRI management compared with transplanted and flooded rice plants, considering also the effects of different means of fertilization and different varieties in rice. Experiments were conducted in 2015 and 2016 under the tropical climate of Nepal's western terai (plains) during both the rainy season (July to November) and the dry season (March to July). The results indicated significantly better performance (P = 0.01) associated with Trichoderma inoculation for both seasons and for both systems of crop management in terms of grain yield and other growth-contributing factors, compared to non-inoculated rice cropping. Relatively higher effects on grain yield were recorded also with organic compared to inorganic fertilization; for unimproved (heirloom) varieties compared with improved varieties; and from SRI vs. conventional flooded crop management. The yield increase with Trichoderma treatments across all trials was 31% higher than in untreated plots (4.9 vs 4.5 mt ha-1). With Trichoderma treatment, yields compared with non-treated plots were 24% higher with organic SRI (6.38 vs 5.13 mt ha-1) and 52% higher with non-organic SRI (6.38 vs 3.53 mt ha-1). With regard to varietal differences, under SRI management Trichoderma inoculation of the improved variety Sukhadhan-3 led to 26% higher yield (6.35 vs 5.04 mt ha-1), and with the heirloom variety Tilkidhan, yield was 41% higher (6.29 vs 4.45 mt ha-1). Economic analysis indicated that expanding the organic cultivation of local landraces under SRI management should be profitable for farmers where such rice has a good market price due to its premium quality and high demand and when SRI enhances yield. These varieties' present low yields can be significantly increased by integrating Trichoderma bio-inoculation with SRI cultural methods. Other recent research has shown that such inoculation can be managed profitably by farmers themselves.

Rhizobia promote the growth of rice shoots by targeting cell signaling, division and expansion.

KEY MESSAGE: The growth-promotion of rice seedling following inoculation with Sinorhizobium meliloti 1021 was a cumulative outcome of elevated expression of genes that function in accelerating cell division and enhancing cell expansion. Various endophytic rhizobacteria promote the growth of cereal crops. To achieve a better understanding of the cellular and molecular bases of beneficial cereal-rhizobia interactions, we performed computer-assisted microscopy and transcriptomic analyses of rice seedling shoots (Oryza sativa) during early stages of endophytic colonization by the plant growth-promoting Sinorhizobium meliloti 1021. Phenotypic analyses revealed that plants inoculated with live rhizobia had increased shoot height and dry weight compared to control plants inoculated with heat-killed cells of the same microbe. At 6 days after inoculation (DAI) with live cells, the fourth-leaf sheaths showed significant cytological differences including their enlargement of parenchyma cells and reduction in shape complexity. Transcriptomic analysis of shoots identified 2,414 differentially-expressed genes (DEGs) at 1, 2, 5 and 8 DAI: 195, 1390, 1025 and 533, respectively. Among these, 46 DEGs encoding cell-cycle functions were up-regulated at least 3 days before the rhizobia ascended from the roots to the shoots, suggesting that rhizobia are engaged in long-distance signaling events during early stages of this plant-microbe interaction. DEGs involved in phytohormone production, photosynthetic efficiency, carbohydrate metabolism, cell division and wall expansion were significantly elevated at 5 and 8 DAI, consistent with the observed phenotypic changes in rice cell morphology and shoot growth-promotion. Correlation analysis identified 104 height-related DEGs and 120 dry-weight-related DEGs that represent known quantitative-trait loci for seedling vigor and increased plant height. These findings provide multiple evidences of plant-microbe interplay that give insight into the growth-promotion processes associated with this rhizobia-rice beneficial association.

Yield effect of applying earthworm castings produced during the oilseed rape-growing season in rice-oilseed rape cropping fields to rice.

In-field earthworm density can be increased by planting oilseed rape during the non-rice growing season as compared to maintaining the rice-growing fields in fallow. This study was conducted to determine the effect on rice yield of earthworm castings produced during the oilseed rape-growing season in rice-oilseed rape cropping fields and to identify the critical factors that contribute to the yield effect. Field microplot experiments were conducted in 2016 and 2017. In 2016, a rice cultivar was grown under a factorial combination of absence (EC0: 0 kg m-2) and presence of earthworm castings (EC1: 17 kg m-2) with three N application rates (9, 12 and 15 g m-2). In 2017, nine rice cultivars were grown under EC0 and EC1 with the moderate N rate as was used in 2016. Results showed that application of earthworm castings produced during the oilseed rape-growing season in rice-oilseed rape cropping fields had a significant positive yield effect on rice. This was attributed to increased panicle m-2 and total aboveground biomass while spikelets panicle-1, spikelet filling percentage, grain weight, and harvest index were not affected. Our study indirectly provides a new evidence that oilseed rape is an excellent previous crop for cereals.

Higher yields of hybrid rice do not depend on nitrogen fertilization under moderate to high soil fertility conditions.

BACKGROUND: Increasing rice yield with fewer external inputs is critical to ensuring food security, reducing environmental costs, and improving returns. Use of hybrid rice has expanded greatly in China due to its higher yield potential. Meanwhile, large and increasing amounts of nitrogen (N) fertilizers have been used for expanding rice production in China. It is not clear to what extent the success of hybrid rice in China is associated with N fertilizer inputs. FINDINGS: We observed that the higher grain yield with N fertilizer in hybrid rice was driven more by a higher yield without N fertilizer than by increases in grain yield with N fertilizer under moderate to high soil fertility conditions. CONCLUSIONS: Our results suggest that greater application of N fertilizers is not needed to benefit from hybrid rice production under moderate to high soil fertility conditions, and that improving and maintaining soil fertility should be a focus for sustaining hybrid rice production. Moreover, our study also indicates that zero-N testing may be a potentially useful tool to develop hybrid rice with high yield and without requirement of greater external N inputs under moderate to high soil fertility conditions.

Effects of Low Temperature Stress on Spikelet-Related Parameters during Anthesis in Indica-Japonica Hybrid Rice.

Poor spikelet fertility under low temperature (LT) stress during anthesis limits the possibility of high yield potential in indica-japonica hybrid rice, leading to reduced stability of grain yield. However, the cause for it is still unclear. In this study, three indica-japonica hybrid rice cultivars, Yongyou9, Yongyou17 (both cold sensitive), and Yongyou538, and one japonica inbred rice cultivar, Zhejing88 (cold tolerant), were grown under LT (17°C) and ambient temperature (AT) (25°C) during anthesis to test for their response with respect to spikelet fertility, pollen germination, and spikelet flowering. The results indicated that LT resulted in lower spikelet fertility in cold-sensitive cultivars than in cold-tolerant cultivars. Spikelet fertility was highly correlated with pollen germination on the stigma. The number of pollen grains and germinated pollen were higher in cold-tolerant cultivars than in cold-sensitive cultivars. Pollen fertility and pollen diameter were also higher in cold-tolerant cultivars, although the latter could achieve a high number of spikelets at anthesis in flowering patterns throughout the duration of LT stress. There were significant differences in anther width and volume between genotypes and treatments according to microscopic analyses, but no differences were observed in anther dehiscence. Moreover, variation in the number of pollen grains on stigmas and in spikelet fertility was not related to either the number of spikelets reaching anthesis or anther dehiscence. Overall, improved anther size, better pollen function, and higher spikelet fertility under LT stress were observed in cold-tolerant cultivars than in cold-sensitive cultivars. The results suggest that the increase in spikelet fertility is due to enhanced pollen germination rather than the number of spikelets reaching anthesis.