Phosphorus deficiency promotes root:shoot ratio and carbon accumulation via modulating sucrose utilization in maize.

Phosphorus deficiency usually promotes root:shoot ratio and sugar accumulation. However, how the allocation and utilization of carbon assimilates are regulated by phosphorus deficiency remains unclear. To understand how phosphorus deficiency affects the allocation and utilization of carbon assimilates, we systematically investigated the fixation and utilization of carbon, along with its diurnal and spatial patterns, in hydroponically grown maize seedlings under low phosphorus treatment. Under low phosphorus, sucrolytic activity was slightly inhibited by 12.0% in the root but dramatically inhibited by 38.8% in the shoot, corresponding to the promoted hexose/sucrose ratio and biomass in the root. Results point to a stable utilization of sucrose in the root facilitating competition for more assimilates, while increasing root:shoot ratio. Moreover, starch and sucrose accumulated in the leaves under low phosphorus. Spatially, starch and sucrose were oppositely distributed, starch mainly in the leaf tip, and sucrose mainly in the leaf base and sheath. Evidence of sucrose getting stuck in leaf base and sheath suggests that carbon accumulation is not attributed to carbon assimilation or export disturbance, but may be due to poor carbon utilization in the sinks. These findings improve the understanding of how low phosphorus regulates carbon allocation between shoot and root for acclimation to stress, and highlight the importance of improving carbon utilization in sinks to deal with phosphorus deficiency.

Multiple cropping effectively increases soil bacterial diversity, community abundance and soil fertility of paddy fields.

BACKGROUND: Crop diversification is considered as an imperative approach for synchronizing the plant nutrient demands and soil nutrient availability. Taking two or more crops from the same field in one year is considered as multiple cropping. It improves the diversity and abundance of soil microbes, thereby improving the growth and yield of crops. Therefore, the present study was conducted to explore the effects of different multiple winter cropping on soil microbial communities in paddy fields. In this study, eight rice cropping patterns from two multiple cropping systems with three different winter crops, including Chinese milk vetch (CMV), rape, and wheat were selected. The effects of different multiple winter cropping on soil microbial abundance, community structure, and diversity in paddy fields were studied by 16 S rRNA high-throughput sequencing and real-time fluorescence quantitative polymerase chain reaction (PCR). RESULTS: The results showed that different multiple winter cropping increased the operational taxonomic units (OTUs), species richness, and community richness index of the bacterial community in 0 ~ 20 cm soil layer. Moreover, soil physical and chemical properties of different multiple cropping patterns also affected the diversity and abundance of microbial bacterial communities. The multiple cropping increased soil potassium and nitrogen content, which significantly affected the diversity and abundance of bacterial communities, and it also increased the overall paddy yield. Moreover, different winter cropping changed the population distribution of microorganisms, and Proteobacteria, Acidobacteria, Nitrospira, and Chloroflexi were identified as the most dominant groups. Multiple winter cropping, especially rape-early rice-late rice (TR) andChinese milk vetch- early rice-late rice (TC) enhanced the abundance of Proteobacteria, Acidobacteria, and Actinobacteria and decreased the relative abundance of Verrucomicrobia and Euryarchaeota. CONCLUSION: In conclusion, winter cropping of Chinese milk vetch and rape were beneficial to improve the soil fertility, bacteria diversity, abundance and rice yield.

Exogenous methylglyoxal alleviates drought-induced 'plant diabetes' and leaf senescence in maize.

Drought-induced leaf senescence is associated with high sugar levels, which bears some resemblance to the syndrome of diabetes in humans; however, the underlying mechanisms of such 'plant diabetes' on carbon imbalance and the corresponding detoxification strategy are not well understood. Here, we investigated the regulatory mechanism of exogenous methylglyoxal (MG) on 'plant diabetes' in maize plants under drought stress applied via foliar spraying during the grain-filling stage. Exogenous MG delayed leaf senescence and promoted photoassimilation, thereby reducing the yield loss induced by drought by 14%. Transcriptome and metabolite analyses revealed that drought increased sugar accumulation in leaves through inhibition of sugar transporters that facilitate phloem loading. This led to disequilibrium of glycolysis and overaccumulation of endogenous MG. Application of exogenous MG up-regulated glycolytic flux and the glyoxalase system that catabolyses endogenous MG and glycation end-products, ultimately alleviating 'plant diabetes'. In addition, the expression of genes facilitating anabolism and catabolism of trehalose-6-phosphate was promoted and suppressed by drought, respectively, and exogenous MG reversed this effect, implying that trehalose-6-phosphate signaling in the mediation of 'plant diabetes'. Furthermore, exogenous MG activated the phenylpropanoid biosynthetic pathway, promoting the production of lignin and phenolic compounds, which are associated with drought tolerance. Overall, our findings indicate that exogenous MG activates defense-related pathways to alleviate the toxicity derived from 'plant diabetes', thereby helping to maintain leaf function and yield production under drought.

Dictamnine suppresses the development of pear ring rot induced by Botryosphaeria dothidea infection by disrupting the chitin biosynthesis.

Ring rot induced by Botryosphaeria dothidea is a major cause of growth and postharvest losses in various fruits. There is an urgent need to develop green fungicides due to pesticide resistance and environmental pressure. Here, we demonstrated the efficacy of dictamnine (DIC, 4-methoxyfuro [2,3-ß] quinoline, purity 98%), a compound isolated from the stems and leaves of Clausena lansium, in effectively suppressing pear ring rot by inhibiting the mycelial growth of B. dothidea. The median effective concentration of DIC was 15.48 µg/mL. Application of DIC to B. dothidea resulted in structural disruption of the cell wall and plasma membrane, leading to mycelial deformation, breakage, and cell death. Transcriptome analysis revealed significant inhibition of the synthetic pathways for fungal cell wall and membrane components by DIC. Particularly, the expression of chitin synthase, a key enzyme of chitin synthesis, was prominently down-regulated. Moreover, the chitin content in DIC-treated B. dothidea mycelia exhibited a substantial dose-dependent reduction. Based on these results, it is promising to develop DIC as an antifungal pesticide for controlling ring rot disease in pear fruits. Our study provides new insights into the underlying mechanism through which DIC inhibits the mycelial growth of B. dothidea.

Coordination of carbon assimilation, allocation, and utilization for systemic improvement of cereal yield.

The growth of yield outputs is dwindling after the first green revolution, which cannot meet the demand for the projected population increase by the mid-century, especially with the constant threat from extreme climates. Cereal yield requires carbon (C) assimilation in the source for subsequent allocation and utilization in the sink. However, whether the source or sink limits yield improvement, a crucial question for strategic orientation in future breeding and cultivation, is still under debate. To narrow the knowledge gap and capture the progress, we focus on maize, rice, and wheat by briefly reviewing recent advances in yield improvement by modulation of i) leaf photosynthesis; ii) primary C allocation, phloem loading, and unloading; iii) C utilization and grain storage; and iv) systemic sugar signals (e.g., trehalose 6-phosphate). We highlight strategies for optimizing C allocation and utilization to coordinate the source-sink relationships and promote yields. Finally, based on the understanding of these physiological mechanisms, we envisage a future scenery of "smart crop" consisting of flexible coordination of plant C economy, with the goal of yield improvement and resilience in the field population of cereals crops.

Inhibition of rice germination by ustiloxin A involves alteration in carbon metabolism and amino acid utilization.

Ustiloxins are the main mycotoxin in rice false smut, a devastating disease caused by Ustilaginoidea virens. A typical phytotoxicity of ustiloxins is strong inhibition of seed germination, but the physiological mechanism is not clear. Here, we show that the inhibition of rice germination by ustiloxin A (UA) is dose-dependent. The sugar availability in UA-treated embryo was lower while the starch residue in endosperm was higher. The transcripts and metabolites responsive to typical UA treatment were investigated. The expression of several SWEET genes responsible for sugar transport in embryo was down-regulated by UA. Glycolysis and pentose phosphate processes in embryo were transcriptionally repressed. Most of the amino acids detected in endosperm and embryo were variously decreased. Ribosomal RNAs for growth were inhibited while the secondary metabolite salicylic acid was also decreased under UA. Hence, we propose that the inhibition of seed germination by UA involves the block of sugar transport from endosperm to embryo, leading to altered carbon metabolism and amino acid utilization in rice plants. Our analysis provides a framework for understanding of the molecular mechanisms of ustiloxins on rice growth and in pathogen infection.

Late Split-Application with Reduced Nitrogen Fertilizer Increases Yield by Mediating Source-Sink Relations during the Grain Filling Stage in Summer Maize.

In the North China Plain, the excessive application of nitrogen (N) fertilizer for ensuring high yield and a single application at sowing for simplifying management in farmer practice lead to low N use efficiency and environmental risk in maize (Zea mays L.) production. However, it is unclear whether and how late split application with a lower level of N fertilizer influences maize yield. To address this question, a two-year field experiment was conducted with two commercial maize cultivars (Zhengdan 958 and Denghai 605) using a lower level of N input (180 kg ha-1) by setting up single application at sowing and split application at sowing and later stages (V12, R1, and R2) with four different ratios, respectively. The maize yield with split-applied 180 kg ha-1 N did not decrease compared to the average yield with 240 kg ha-1 N input in farmer practice, while it increased by 6.7% to 11.5% in the four N split-application treatments compared with that of the single-application control. Morphological and physiological analyses demonstrated that late split application of N (i) increased the net photosynthetic rate and chlorophyll content and thus promoted the photosynthetic efficiency during the reproductive stages; (ii) promoted the sink capacity via improved kernel number, endosperm cells division, and grain-filling rate; and (iii) increased the final N content and N efficiency in the plant. Therefore, we propose that late split application of N could reduce N fertilizer input and coordinately improve N efficiency and grain yield in summer maize production, which are likely achieved by optimizing the source-sink relations during the grain-filling stage.

A transcriptional landscape underlying sugar import for grain set in maize.

Developing seed depends on sugar supply for its growth and yield formation. Maize (Zea mays L.) produces the largest grains among cereals. However, there is a lack of holistic understanding of the transcriptional landscape of genes controlling sucrose transport to, and utilization within, maize grains. By performing in-depth data mining of spatio-temporal transcriptomes coupled with histological and heterologous functional analyses, we identified transporter genes specifically expressed in the maternal-filial interface, including (i) ZmSWEET11/13b in the placento-chalazal zone, where sucrose is exported into the apoplasmic space, and (ii) ZmSTP3, ZmSWEET3a/4c (monosaccharide transporters), ZmSUT1, and ZmSWEET11/13a (sucrose transporters) in the basal endosperm transfer cells for retrieval of apoplasmic sucrose or hexoses after hydrolysis by extracellular invertase. In the embryo and its surrounding regions, an embryo-localized ZmSUT4 and a cohort of ZmSWEETs were specifically expressed. Interestingly, drought repressed those ZmSWEETs likely exporting sucrose but enhanced the expression of most transporter genes for uptake of apoplasmic sugars. Importantly, this drought-induced fluctuation in gene expression was largely attenuated by an increased C supply via controlled pollination, indicating that the altered gene expression is conditioned by C availability. Based on the analyses above, we proposed a holistic model on the spatio-temporal expression of genes that likely govern sugar transport and utilization across maize maternal and endosperm and embryo tissues during the critical stage of grain set. Collectively, the findings represent an advancement towards a holistic understanding of the transcriptional landscape underlying post-phloem sugar transport in maize grain and indicate that the drought-induced changes in gene expression are attributable to low C status.

Differential ear growth of two maize varieties to shading in the field environment: Effects on whole plant carbon allocation and sugar starvation response.

The interception of irradiation by smog pollution and cloud cover associated with extreme rainfall events has become an increasingly important limiting factor in crop production in China. Little is known about the adaptation of carbon (C) allocation to periodic low irradiance in field conditions. The trehalose signaling pathway plays a critical role in adapting C allocation to the environment in crops but its importance in adaptation to low light in field conditions is not known. To determine the effects of low irradiance on C economy and maize yield, two commonly grown hybrids (LY-16 and ZD-958) were subject to three levels of shading (15 %, 50 %, and 97 %) for one week from V13 stage in two successive seasons. Shading led to yield loss mainly due to decreased kernel number, which was greater in LY-16 than ZD-958. Effects of shading on leaf area and photosynthesis were similar in both varieties. Starch levels in leaves were maintained, whereas total soluble carbohydrates were reduced up to fivefold by shading in both varieties. Shading increased the proportion of photoassimilate retained in leaves relative to reproductive organs. Carbohydrates in ears and stem were decreased by shading similarly in both varieties. Amongst the parameters measured, the main difference between LY-16 and ZD-958 associated with yield penalty was the expression of class II trehalose phosphate synthase (TPS) and trehalose-6-phosphate phosphatase (TPP) genes which were increased due to shading in leaves and ears, particularly in ears of LY-16. It is concluded that altered C fixation and allocation by low irradiance limited ear growth at pre-anthesis. Activation of TPSII and TPP genes indicates that the trehalose pathway likely plays a role in ear development under low light and could be a target for yield improvement under such conditions as with other stresses.

Intervening in sibling competition for assimilates by controlled pollination prevents seed abortion under postpollination drought in maize.

During maize production, drought throughout the flowering stage usually induces seed abortion and yield losses. The influence of postpollination drought stress on seed abortion and its underlying mechanisms are not well characterized. By intervening in the competition for assimilates between kernel siblings under different degrees of postpollination drought stresses accompanied by synchronous pollination (SP) and incomplete pollination (ICP) approaches, the mechanisms of postpollination abortion were investigated at physiological and molecular levels. Upon SP treatment, up to 15% of the fertilized apical kernels were aborted in the drought-exacerbated competition for assimilates. The aborted kernels exhibited weak sucrose hydrolysis and starch synthesis but promoted the synthesis of trehalose-6-phosphate and ethylene. In ICP where basal pollination was prevented, apical kernel growth was restored with reinstated sucrose metabolism and starch synthesis and promoted sucrose and hexose levels under drought stress. In addition, the equilibrium between ethylene and polyamine in response to the drought and pollination treatments was associated with the abortion process. We conclude that competition for assimilates drives postpollination kernel abortion, whereas differences in sugar metabolism and the equilibrium between ethylene and polyamines may be relevant to the "live or die" choice of kernel siblings during this competition.

Seasonal and diurnal patterns of non-structural carbohydrates in source and sink tissues in field maize.

BACKGROUND: Carbohydrate partitioning and utilization is a key determinant of growth rate and of yield in plants and crops. There are few studies on crops in field conditions. In Arabidopsis, starch accumulation in leaves is a negative indicator of growth rate. RESULTS: Here, we wished to establish if starch accumulation in leaves could potentially be a marker for growth rate and yield in crops such as maize. We characterized daily patterns of non-structural carbohydrate (NSC) at different growth stages over two seasons for maize hybrids in the field. In 27 commercial hybrids, we found a significant negative relationship between residual starch in leaves and plant growth, but not with final yield and biomass. We then focused on three typical hybrids and established a method for calculation of C turnover in photosynthetic leaves that took into account photosynthesis, leaf area and NSC accumulation. The ratios of stored NSC decreased from approximately 15% to less than 4% with ongoing ontogeny changes from V7 to 28 days after pollination. CONCLUSION: The proportion rather than absolute amount of carbon partitioned to starch in leaves at all stages of development related well with yield and biomass accumulation. It is proposed that screening plants at an early vegetative growth stage such as V7 for partitioning into storage may provide a prospective method for maize hybrid selection. Our study provides the basis for further validation as a screening tool for yield.

Delayed pollination and low availability of assimilates are major factors causing maize kernel abortion.

Selective seed abortion is a survival strategy adopted by many species that sacrifices some seeds to allow the remaining ones to set. While in evolutionary terms this is a successful approach, it causes huge losses to crop yields. A pollination time gap (PTG) has been suggested to be associated with position-related grain abortion. To test this hypothesis, we developed a novel approach to alter the natural pattern of maize (Zea mays L.) pollination and to examine the impact of PTGs on kernel growth and the underlying physiological basis. When apical and basal kernels were synchronously pollinated, the basal kernels set and matured but the apical kernels were aborted at an early stage. Delaying pollination to the basal ovaries suppressed their development and reduced invertase activity and sugar levels, which allowed the apical kernels to set and grow normally. In situ localization revealed normal cell wall invertase activity in apical and basal kernels under synchronous pollination but reduced activity in the delayed-pollinated kernels independent of their position. Starch, which was abundant in basal kernel areas, was absent in the apical kernel regions under synchronous pollination but apparent with delayed pollination. Our analyses identified PTG-related sink strength and a low level of local assimilates as the main causes of grain abortion.

Regulation of maize kernel weight and carbohydrate metabolism by abscisic acid applied at the early and middle post-pollination stages in vitro.

Abscisic acid (ABA) accumulates in plants under drought stress, but views on the role of ABA in kernel formation and abortion are not unified. The response of the developing maize kernel to exogenous ABA was investigated by excising kernels from cob sections at four days after pollination and culturing in vitro with different concentrations of ABA (0, 5, 10, 100µM). When ABA was applied at the early post-pollination stage (EPPS), significant weight loss was observed at high ABA concentration (100µM), which could be attributed to jointly affected sink capacity and activity. Endosperm cells and starch granules were decreased significantly with high concentration, and ABA inhibited the activities of soluble acid invertase and acid cell wall invertase, together with earlier attainment of peak values. When ABA was applied at the middle post-pollination stage (MPPS), kernel weight was observably reduced with high concentration and mildly increased with low concentration, which was regulated due to sink activity. The inhibitory effect of high concentration and the mild stimulatory effect of low concentration on sucrose synthase and starch synthase activities were noted, but a peak level of ADP-glucose pyrophosphorylase (AGPase) was stimulated in all ABA treatments. Interestingly, AGPase peak values were advanced by low concentration and postponed by high concentration. In addition, compared with the control, the weight of low ABA concentration treatments were not statistically significant at the two stages, whereas weight loss from high concentration applied at EPPS was considerably obvious compared with that of the MPPS, but neither led to kernel abortion. The temporal- and dose-dependent impacts of ABA reveal a complex process of maize kernel growth and development.