Molecular mechanisms of stress resistance in sorghum: Implications for crop improvement strategies

Abiotic stresses, such as drought, salt, extreme temperatures, and heavy metal pollution, are the main environmental factors that limit crop growth and yield. Sorghum, a C4 grass plant with high photosynthetic efficiency, can grow in adverse environmental conditions due to its excellent stress resistance characteristics. Therefore, unraveling the stress-resistance mechanism of sorghum could provide a theoretical basis for developing and cultivating various stress-resistant crops. This understanding could also help to create a conducive environment for using marginal soil in agriculture and ensuring food security. In this review, we discuss the adaptation mechanisms of sorghum under drought, salinity, temperature, and soil heavy metal stresses, the specific response to stress, the screening of sorghum-resistant germplasm, and the identification and functional analysis of the relevant genes and quantitative trait loci(QTL). In addition, we discuss the application potential of different stress-tolerant sorghum germplasms reported to date and emphasize the feasibility and potential use in developing and promoting highly stress-tolerant sorghum in marginal soil.

干旱胁迫对2种光合类型C_4荒漠植物叶片光合特征酶和抗氧化酶活性的影响

以荒漠C4草本植物蔷薇猪毛菜(NADP苹果酸酶型,NADP-ME)和粗枝猪毛菜(NAD苹果酸酶型,NADME)为研究对象,采用盆栽控水试验设置正常供水和轻度、中度、重度干旱处理(土壤含水量分别为田间持水量80%、60%、45%和35%),通过测定不同程度干旱胁迫下叶片含水量、C4光合特征酶和抗氧化酶活性等指标,探讨不同类型C4荒漠植物光合特征酶和抗氧化系统对干旱逆境的适应机制。结果显示:(1)2种植物叶片含水量均随干旱胁迫的加剧不同程度降低。(2)叶片磷酸烯醇式丙酮酸羧化酶(PEPC)活性在中度干旱胁迫下显著增加而在重度干旱胁迫下急剧下降;蔷薇猪毛菜NAD-ME活性和粗枝猪毛菜NADP-ME活性都很低,且它们基本不受干旱胁迫的影响;随干旱胁迫的加剧,蔷薇猪毛菜NADP-ME活性呈下降趋势,而粗枝猪毛菜NAD-ME活性先显著增加而在重度干旱胁迫下显著降低。(3)随着干旱胁迫的加剧,叶片超氧化物歧化酶(SOD)活性呈下降趋势,过氧化物酶(POD)活性在不同程度干旱胁迫下均有不同程度增加;过氧化氢酶(CAT)活性在中度干旱胁迫下均有不同程度的增加,但在重度干旱胁迫下蔷薇猪毛菜CAT活性降低,而粗枝猪毛菜CAT活性显著增加;丙二醛(MDA)含量随干旱胁迫的加剧均有不同程度的增加。研究认为,一定程度干旱胁迫下,2种荒漠植物的PEPC活性均有增加;不同光合类型C4植物叶片脱羧酶(NADP-ME和NAD-ME)对干旱胁迫的响应有明显的差异。POD和CAT是这两种C4植物适应干旱胁迫的主要抗氧化酶,但蔷薇猪毛菜CAT在重度干旱胁迫下没有起到积极保护作用。

Simulating leaf net CO_2 assimilation rate of C_3 & C_4 plants and its response to environmental factors

Basic structure and algorithm of leaf mechanism photosynthesis model were described in first part of this study based on former researcher results. Then, considering some environmental factors influencing on leaf photosynthesis, three numerical sensitivity experiments were carried out. We simulated the sing le leaf net CO2 assimilation, which acts as a function of different light, carbo n dioxide and temperature conditions. The relationships between leaf net photosy nthetic rate of C3 and C4 plant with CO2 concentration intercellular, leaf tempe rature, and photosynthetic active radiation (PAR) were presented, respectively. The results show the numerical experiment may indicate the main characteristic o f plant photosynthesis in C3 and C4 plant, and further can be used to integrate with the regional climate model and act as land surface process scheme, and bett er understand the interaction between vegetation and atmosphere.

基于生物质能的芒属(Miscanthus)植物碳动态和收支研究进展

生物质能(Biomass energy)是最为广泛的可再生能源,其中多年生芒属C4植物(Miscanthus)由于具有巨大碳固定能力而成为潜力巨大的生物质能植物。中国是芒属植物芒草起源中心,但相对于欧洲国家应对能源危机和温室效应而采取的芒草研究与应用来说,仍处于起步阶段。我国长期以来传统的草地利用模式,决定了在南方草地的研究显著少于北方,近年来芒草在华南地区的运用研究集中于生态修复,对草本植物群落基于生态系统水平的CO2气体交换能力的研究仍然相当缺乏,在二氧化碳浓度持续增长及全球变暖背景下,生物质能植物及其碳汇功能的相关研究尤显重要。我国南方近6 700万hm2退化丘陵草坡急待恢复或处于恢复中,草坡地芒属植物符合生物质能植物标准,施肥少,害虫少,农药输入少,能够有效地利用光、水等自然资源。考虑到C4植物具有比C3植物更强的光合作用能力,高光能利用率C4芒属植物的碳固定能力及能源潜力值得重视,但缺乏科学的碳动态和碳收支评估。综述了国内外芒草生物量特征与生物质能潜力研究现状,重点论述芒属植物生态系统水平的碳动态和收支能力研究,探讨了系统水平更客观评估芒属碳源汇(Carbon sequestration)功能的方法,基于生物量过程的研究结果及华南地区草坡研究历史和现状,为草坡地生物质能的合理开发利用提出了相关对策,强调在我国南方开发和利用芒属植物资源具有重要能源价值和经济、环境效益。

植物丙酮酸磷酸双激酶研究进展

丙酮酸磷酸双激酶(pyruvate orthophosphate dikinase,PPDK)作为C4光合途径中一个非常重要的限速酶,其功能已经清楚,但在C3植物中以及逆境条件下的作用尚不明确。在阐述PPDK基本生物学特征的基础上,重点介绍了PPDK在C4植物和C3植物中的功能、活性调控、基因工程以及PPDK对逆境胁迫应答的研究进展,以期为植物抗逆基因挖掘及抗逆种质创制提供参考。

Simulating leaf net CO2 assimilation rate of C3 & C4 plants and its response to environmental factors

Basic structure and algorithm of leaf mechanism photosynthesis model were described in first part of this study based on former researcher results. Then, considering some environmental factors influencing on leaf photosynthesis, three numerical sensitivity experiments were carried out. We simulated the sing le leaf net CO2 assimilation, which acts as a function of different light, carbo n dioxide and temperature conditions. The relationships between leaf net photosy nthetic rate of C3 and C4 plant with CO2 concentration intercellular, leaf tempe rature, and photosynthetic active radiation (PAR) were presented, respectively. The results show the numerical experiment may indicate the main characteristic o f plant photosynthesis in C3 and C4 plant, and further can be used to integrate with the regional climate model and act as land surface process scheme, and bett er understand the interaction between vegetation and atmosphere.

Response of root traits of Reaumuria soongorica and Salsola passerina to facilitation

C3 plant Reaumuria soongorica and C4 plant Salsola passerina are super xerophytes and coexist in a mixed community in either isolated or associated growth, and interspecific facilitation occurs in associated growth. In the present study, the root traits including root distribution, root length（RL）, root surface area（RSA）, root weight（RW） and specific root length（SRL） of both species in two growth forms were investigated to clarify their response to facilitation in associated growth. Six isolated plants of each species, as well as six associated plants similar in size and development were selected during the plant growing season, and their roots were excavated at 0–10, 10–20, 20–30, 30–40 and 40–50 cm soil depths at the end of the growing season. All the roots of each plant were separated into the two categories of fine roots（2 mm diameter） and coarse roots（≥2 mm diameter）. Root traits such as RL and RSA in the fine and coarse roots were obtained by the root analyzing system WinRHIZO. Most of the coarse roots in R. soongorica and S. passerina were distributed in the top 10 cm of the soil in both growth forms, whereas the fine roots of the two plant species were found mainly in the 10–20 and 20–30 cm soil depths in isolated growth, respectively. However, the fine roots of both species were mostly overlapped in 10–20 cm soil depth in associated growth. The root/canopy ratios of both species reduced, whereas the ratios of their fine roots to coarse roots in RL increased, and both species had an increased SRL in the fine roots in associated growth. In addition, there was the increase in RL of fine roots and content of root N for S. passerina in associated growth. Taken together, the root growth of S. passerina was facilitated for water and nutrient exploration under the interaction of the overlapped roots in both species in associated growth, and higher SRL allowed both species to more effectively adapt to the infertile soil in the desert ecosystem.

Carbon isotope variations of modern soils in the eastern margin of the Tibetan Plateau and their controlling factors

The marginal areas of the Tibetan Plateau have great vertical altitude gradient and abundant vegetation, they are therefore the ideal places for investigating the relationships among carbon isotope composition(δ^(13)C) of modern soils, vegetation and environmental factors, which would be very useful for the reconstructions of both paleovegetation and paleoclimate. In this paper, modern soil samples collected in different vegetation vertical zones along 4km elevation gradient in the eastern margin of the Tibetan Plateau were analyzed for their carbon isotope composition. The results show that the modern soils in different vegetation vertical zones show apparent difference of δ^(13)C values, which get heavier in the sequence of mixed evergreen and deciduous broad-leaved forest(-27.28‰ on average), evergreen broad-leaved forest(-27.25‰), subalpine shrub-meadow(-25.81‰), subalpine coniferous forest(-25.81‰), alpine bush-meadow(-25.16‰), and drought-enduring shrub(-24.07‰). 1800 m and 3500 m are two critical points for the δ^(13)C values with respect to altitude. Specifically, the δ^(13)C values decrease with increasing altitude below both points while increasing with increasing altitude above both points. Further analyses indicate that the declining δ^(13)C values are mainly controlled by the decreasing proportion of C4 plants with elevation and the increasing δ^(13)C values are attributed to the plant physic-morphological adaptation to the alpine environment. In the absence of drought stress, temperature is the main controlling factor for the carbon isotopic variations with altitude gradient.

30种藜科C_4植物同化器官解剖结构及δ^(13)C值

观测了藜科滨藜族(Atripliceae)、樟味藜族(Camphorosmeae)、碱蓬族(Suaedeae)和猪毛菜族(Salsoleae)30种C4植物的同化器官的解剖结构和δ13 C值,共见到12种解剖结构类型。滨藜族包含2种类型,Atriplicoid-Ⅰ型和Atriplicoid-Ⅱ型,它们的差异仅在于结构中是否含有皮下组织。樟味藜族包括3种解剖结构类型,即含有皮下组织的Kochioid-Ⅰ型、叶肉细胞和维管束鞘细胞形成近似于环形的Kochioid-Ⅱ型和不含有皮下组织的Kochioid-Ⅲ型。碱蓬族包含3种类型,Kranz-Suaedoid-Ⅰ型和Kranz-Suaedoid-Ⅱ型结构中具有皮下组织,但是Kranz-Suaedoid-Ⅱ型结构中只具有一层叶肉细胞,这种结构存在于异子蓬属的异子蓬(Borszczowia aralocaspica),而Kranz-Suaedoid-Ⅲ型结构中不具有皮下组织。猪毛菜族植物叶片/同化枝的解剖结构类型有4种,它们的差异在于结构中是否含有皮下组织、次要的较小的维管束的位置以及环形的维管束鞘细胞是否被打断。30种植物的δ13 C值范围-11.1‰^-14.5‰,显示为C4植物,与其同化器官解剖结构相对应。

C_4植物玉米的光合-光响应曲线模拟研究

光是植物光合作用最基本的一个决定因子,准确分析光响应曲线及其参数是研究光合生理生态过程对环境变化响应的重要途径;但相关模型及其模拟的准确性仍待改进。该研究基于C_4作物玉米(Zea mays)不同干旱处理试验资料,比较研究了现有光响应模型(直角双曲线模型、非直角双曲线模型、直角双曲线修正模型、指数模型、二次函数模型以及新提出的改进模型)的适应性。结果表明,改进的光响应模型具有较好的精确度,可较准确地描述光响应曲线,也能够准确拟合最大净光合速率、光饱和点、光补偿点以及暗呼吸速率4个关键光合参数。该结果为研究植物光合生理生态过程及其环境适应性提供了一个改进的模拟方法。

Hybrid Rubisco with Complete Replacement of Rice Rubisco Small Subunits by Sorghum Counterparts Confers C4 Plant-like High Catalytic Activity

Photosynthetic rate at the present atmospheric condition is limited by the CO2-fixing enzyme ribulose-1,5-bisphosphate carboxylase/oxygenase(Rubisco)because of its extremely low catalytic rate(kcat)and poor affinity for C02(Kc)and specificity for CO2(Sc/O).Rubisco in C4 plants generally shows higher k cat than that in C3 plants.Rubisco consists of eight large subunits and eight small subunits(RbcS).Previously,the chimeric incorporation of sorghum C4-type RbcS significantly increased the k cat of Rubisco in a C3 plant,rice.In this study,we knocked out rice RbcS multigene family using the CRISPR-Cas9 technology and completely replaced rice RbcS with sorghum RbcS in rice Rubisco.Obtained hybrid Rubisco showed almost C4 plant-like catalytic properties,i.e.,higher kcai,higher Kc,and lower Sc/O.Transgenic lines expressing the hybrid Rubisco accumulated reduced levels of Rubisco,whereas they showed slightly but significantly higher photosynthetic capacity and similar biomass production under high C 02 condition compared with wild-type rice.High-resolution crystal structural analysis of the wild-type Rubisco and hybrid Rubisco revealed the structural differences around the central pore of Rubisco and the pC-pD hairpin in RbcS.We propose that such differences,particularly in the pC-pD hairpin,may impact the flexibility of Rubisco catalytic site and change its catalytic properties.

Contribution of the Alternative Respiratory Pathway to PSII Photoprotection in C3 and C4 Plants

The mechanism by which the mitochondrial alternative oxidase （AOX） pathway contributes to photosystem II （PSII） photoprotection is in dispute. It was generally thought that the AOX pathway protects photosystems by dissipating excess reducing equivalents exported from chloroplasts through the malate/oxaloacetate （Mal/OAA） shuttle and thus preventing the over-reduction of chloroplasts. In this study, using the aoxla Arabidopsis mutant and nine other C3 and C4 plant species, we revealed an additional action model of the AOX pathway in PSII photoprotection. Although the AOX pathway contributes to PSII photoprotection in C3 leaves treated with high light, this contribution was observed to disappear when photorespiration was suppressed. Disruption or inhibition of the AOX pathway significantly decreased the photorespiration in C3 leaves. Moreover, the AOX pathway did not respond to high light and contributed little to PSII photoprotection in C4 leaves possessing a highly active Mal/OAA shuttle but with little photorespiration. These results demonstrate that the AOX pathway contributes to PSII photoprotection in C3 plants by maintaining photo- respiration to detoxify glycolate and via the indirect export of excess reducing equivalents from chloro-plasts by the MaI/OAA shuttle. This new action model explains why the AOX pathway does not contribute to PSII photoprotection in C4 plants.

C_4 Plants as Biofuel Feedstocks: Optimising Biomass Production and Feedstock Quality from a Lignocellulosic Perspective

The main feedstocks for bioethanol are sugarcane （Saccharum offic- inarum） and maize （Zea mays）, both of which are C4 grasses, highly efficient at converting solar energy into chemical energy, and both are food crops. As the systems for lignocellulosic bioethanol production become more efficient and cost effective, plant biomass from any source may be used as a feedstock for bioethanol production. Thus, a move away from using food plants to make fuel is possible, and sources of biomass such as wood from forestry and plant waste from cropping may be used. However, the bioethanol industry will need a continuous and reliable supply of biomass that can be produced at a low cost and with minimal use of water, fertilizer and arable land. As many C4 plants have high light, water and nitrogen use efficiency, as compared with C3 species, they are ideal as feedstock crops. We consider the productivity and resource use of a number of candidate plant species, and discuss biomass ＇quality＇, that is, the composition of the plant cell wall.

Investigation of the controlled factors influencing carbon isotope composition of foxtail and common millet on the Chinese Loess Plateau

To apply carbon isotope composition （δ13C） analyses of C4 plants to the quantitative reconstruction of paleoclimate, the functional mechanism linking plant δ13C （δ13Cp） to the environment, which is based on the plants＇ physiological characteristics and morphological adaptability, must be thoroughly understood. Foxtail millet （Setaria italic） and common millet （Panicum miliaceum）, as C4 plants, are representative crops of the rain-fed agriculture present in northern China. Fossil millets are ideal for paleoclimatic studies because of the ease of acquisition and identification to the species level. Modem seeds of foxtail and common millet collected from different habitats of the Chinese Loess Plateau, and their carbon isotope compositions, were an- alyzed and correlated with environmental factors, such as latitude, altitude, temperature, precipitation, water availability, and relative humidity. The results showed that the δ13C of foxtail millet had a significantly negative correlation with latitude （R=-0.46）, which may indicate the influence of light. The effect of light on the δ13C of foxtail millet accounted for only 21% of variability, while other climatic factors did not exert significant influences. Thus, the δ13C of foxtail millet was not suitable for extracting climatic information. The δ13C of common millet was significantly and positively correlated with precipitation during the growing period （R=0.75）, explaining 56% of variability. The functional mechanisms analyzed, using the plants＇ physiological characteristics and morphological adaptability, indicated that common millet can adapt to environmental changes because of stomatal sensitivity and some non-stomatal factors. Therefore, the 813C of common millet can record precipitation during growth and is a promising factor for paleoclimatic reconstruction.

CO_(2)emission and source partitioning from carbonate and non-carbonate soils during incubation

The accurate quantification and source partitioning of CO_(2)emitted from carbonate(i.e.,Haplustalf)and non-carbonate(i.e.,Hapludult)soils are critically important for understanding terrestrial carbon(C)cycling.The two main methods to capture CO_(2)released from soils are the alkali trap method and the direct gas sampling method.A 25-d laboratory incubation experiment was conducted to compare the efficacies of these two methods to analyze CO_(2)emissions from the non-carbonate and carbonate-rich soils.An isotopic fraction was introduced into the calculations to determine the impacts on partitioning of the sources of CO_(2)into soil organic carbon(SOC)and soil inorganic carbon(SIC)and into C3 and/or C4 plant-derived SOC.The results indicated that CO_(2)emissions from the non-carbonate soil measured using the alkali trap and gas sampling methods were not significantly different.For the carbonate-rich soil,the CO_(2)emission measured using the alkali trap method was significantly higher than that measured using the gas sampling method from the 14 th day of incubation onwards.Although SOC and SIC each accounted for about 50%of total soil C in the carbonate-rich soil,SOC decomposition contributed 57%–72%of the total CO_(2)emitted.For both non-carbonate and carbonate-rich soils,the SOC derived from C4 plants decomposed faster than that originated from C3 plants.We propose that for carbonate soil,CO_(2)emission may be overestimated using the alkali trap method because of decreasing CO_(2)pressure within the incubation jar,but underestimated using the direct gas sampling method.The gas sampling interval and ambient air may be important sources of error,and steps should be taken to mitigate errors related to these factors in soil incubation and CO_(2)quantification studies.

The precipitation “threshold value” on C_4/C_3 abundance of the Loess Plateau, China

In this paper, we report organic carbon isotopic characterizations from two loess sequences （JY and GL）, spanning the last 20 ka, from the northwest Chinese Loess Plateau （CLP）. The results indicate that the vegetation type is nearly pure C3 plants in the studied region during the Holocene. In contrast to other reported loess sequences in the central-southeast CLP, the relative abundance of the C4 plants decreases from southeast to northwest, with the vegetation types changing from pure C3 to a C4/C3 mixture should near 36°N in the Holocene. From the perspective of the modem temperature and precipitation distribution, the summer temperature has no obvious change at the same latitude, but there are differences in the summer precipi- tation, which exhibit an obvious increase from west to east. Further analysis indicated that the C4 plant abundance decreases with the decreasing summer season precipitation from the southeast to the northwest CLP during the Holocene. We suggest that with the absence of favorable precipitation condition, increasing temperature and decreasing atmospheric CO2 concentration are insufficient to drive an expansion of the C4 plants on the CLP in the Holocene. According to a Holocene precipitation recon- struction, a ＂threshold value＂ of summer precipitation existed, which mainly controlled the expansion of C4 plants. Compared with the modern δ13Csom and climate data on the CLP, both the Holocene and the present ＂threshold value＂ are near the 360-mm summer precipitation line, although the present precipitation line turned slightly southwestward. Our results provide new insights for further research on the C4/C3 variations with precipi- tation and the relationship to global C4/C3 change.

Climate and anthropogenic drivers of changes in abundance of C4 annuals and perennials in grasslands on the Mongolian Plateau

Background:C4 plants have increased substantially during the past several decades in the grasslands of the Mongolian Plateau due to regional warming.Here,we explore how the patterns of abundances of C4 annuals and C4 perennials change over space and time.Methods:A total of 280 sites with C4 plants were surveyed in four types of grasslands in 9 years.The relative biomasses of C4 plants(PC4),C4 annuals(PA4),and C4 perennials(PP4)were calculated.Structural equation modeling was used to analyze the drivers of changes in PA4 and PP4.Results:At the regional scale,PA4 on average was 11%(±19%,SD)and PP4 was 13%(±19%,SD).Spatially,C4 annuals dominated the C4 communities within an east–west belt region along 44°N and tended to spread toward northern latitudes(about 0.5°)and higher altitudes in the east mountainous areas.The abundance of C4 annuals decreased,while that of C4 perennials increased.The patterns of C4 annuals and C4 perennials were mainly controlled by temperature,growing season precipitation,and dynamics between the two life forms.Conclusions:C4 annuals exhibited competitive advantages in normal and wet years,while C4 perennials had competitive advantages in dry years.Grazing as a main human disturbance increased C4 annuals,but had no significant effect on C4 perennials.

Recent progress in the single-cell C4 photosynthesis in terrestrial plants

Currently, single-cell C4 photosynthesis has been reported in four terrestrial plant species, Bienertia cycloptera, B. sinuspersici, B. kavirense and Suaeda aralocaspica, of family Chenopodiaceae. These species possess novel mechanisms of C4 photosynthesis through spatial partitioning of organelles and key enzymes in distinct cytoplasmic domains within single chlorenchyma cells. Anatomical and biochemical studies have shown that the three Bienertia species and S. aralocaspica utilize biochemical and organellar compartmentation to achieve the equivalent spatial separation of Kranz anatomy but within a single photosynthetic cell. These discoveries have challenged the paradigm for C4 photosynthesis in terrestrial plants which had suggested for more than 40 years that the Kranz feature was indispensably required for its C4 function. In this review, we focus on the recent progress in understanding the cellular and molecular mechanisms that control the spatial relationship of organelles in these unique single-cell C4 systems. The demonstrated interaction of dimorphic chloroplasts with microtubules and actin filaments has shed light on the importance of these cytoskeleton components in the intracellular partitioning of organelles. Future perspectives on the potential function of the cytoskeleton in targeting gene products to specific subcellular compartments are discussed.