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.

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.

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.

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.

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.

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.

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.

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.