Photosynthetic Types Research Articles

The root strategy of the C4 grasses tends to be ‘do-it-yourself’

Root resource acquisition strategies play a crucial role in understanding plant water uptake and drought adaptation. However, the interrelationships among mycorrhizal associations, root hair development, and fine root strategies, as well as the disparities between C3 and C4 grasses, remain largely unknown. A pot experiment was conducted to determine leaf gas exchange, root morphology, root hair, mycorrhizal fungi, and biomass allocation of three C4 grasses and four C3 grasses, common species of grasslands in Northeast China, under the control and drought conditions. Compared to the C3 grasses, the C4 grasses increased specific surface area by decreasing tissue density, yet exhibited root hair factor at only 21 % of the C3 grasses. Under the drought conditions, the C4 grasses exhibited more intense and extensive adjustments in root traits, characterized by shifts toward a more conservative morphology with increased root diameter and tissue density, as well as reduced mycorrhizal colonization rates. These adaptations led to a decrease in root absorptive function, which was compensated in the C4 grasses by greater root biomass partitioning and root hair factor. Variances in root strategies between plants functional groups were closely related to leaf photosynthetic rate, water and nitrogen use efficiency. We observed that the C4 grasses prefer direct acquisition of soil resources through the fine root pathway over the root hair or mycorrhizal pathway, suggesting a ‘do-it-yourself’ approach. These findings provide valuable insights into how plant communities of different photosynthetic types might respond to future climate change.

Identifying genomic regions associated with C4 photosynthetic activity and leaf anatomy in Alloteropsis semialata.

C4 photosynthesis is a complex trait requiring multiple developmental and metabolic alterations. Despite this complexity, it has independently evolved over 60 times. However, our understanding of the transition to C4 is complicated by the fact that variation in photosynthetic type is usually segregated between species that diverged a long time ago. Here, we perform a genome-wide association study (GWAS) using the grass Alloteropsis semialata, the only known species to have C3, intermediate, and C4 accessions that recently diverged. We aimed to identify genomic regions associated with the strength of the C4 cycle (measured using δ13C), and the development of C4 leaf anatomy. Genomic regions correlated with δ13C include regulators of C4 decarboxylation enzymes (RIPK), nonphotochemical quenching (SOQ1), and the development of Kranz anatomy (SCARECROW-LIKE). Regions associated with the development of C4 leaf anatomy in the intermediate individuals contain additional leaf anatomy regulators, including those responsible for vein patterning (GSL8) and meristem determinacy (GIF1). The parallel recruitment of paralogous leaf anatomy regulators between A. semialata and other C4 lineages implies the co-option of these genes is context-dependent, which likely has implications for the engineering of the C4 trait into C3 species.

Increased α-ketoglutarate links the C3-C4 intermediate state to C4 photosynthesis in the genus Flaveria.

As a complex trait, C4 photosynthesis has multiple independent origins in evolution. Phylogenetic evidence and theoretical analysis suggest that C2 photosynthesis, which is driven by glycine decarboxylation in the bundle sheath cell, may function as a bridge from C3 to C4 photosynthesis. However, the exact molecular mechanism underlying the transition between C2 photosynthesis to C4 photosynthesis remains elusive. Here, we provide evidence suggesting a role of higher α-ketoglutarate (AKG) concentration during this transition. Metabolomic data of 12 Flaveria species, including multiple photosynthetic types, show that AKG concentration initially increased in the C3-C4 intermediate with a further increase in C4 species. Petiole feeding of AKG increases the concentrations of C4-related metabolites in C3-C4 and C4 species but not the activity of C4-related enzymes. Sequence analysis shows that glutamate synthase (Fd-GOGAT), which catalyzes the generation of glutamate using AKG, was under strong positive selection during the evolution of C4 photosynthesis. Simulations with a constraint-based model for C3-C4 intermediate further show that decreasing the activity of Fd-GOGAT facilitated the transition from a C2-dominant to a C4-dominant CO2 concentrating mechanism. All these results provide insight into the mechanistic switch from C3-C4 intermediate to C4 photosynthesis.

The first released available genome of the common ice plant (Mesembryanthemum crystallinum L.) extended the research region on salt tolerance, C3-CAM photosynthetic conversion, and halophilism

Background The common ice plant (Mesembryanthemum crystallinum L.) is an annual herb belonging to the genus Mesembryanthemum of the family Aizoaceae, native to Southern Africa. Methods We performed shotgun genome paired-end sequencing using the Illumina platform to determine the genome sequence of the ice plants. We assembled the whole genome sequences using the genome assembler “ALGA” and “Redundans”, then released them as available genomic information. Finally, we mainly estimated the potential genomic function by the homology search method. Results A draft genome was generated with a total length of 286 Mb corresponding to 79.2% of the estimated genome size (361 Mb), consisting of 49,782 contigs. It encompassed 93.49% of the genes of terrestrial higher plants, 99.5% of the ice plant transcriptome, and 100% of known DNA sequences. In addition, 110.9 Mb (38.8%) of repetitive sequences and untranslated regions, 971 tRNA, and 100 miRNA loci were identified, and their effects on stress tolerance and photosynthesis were investigated. Molecular phylogenetic analysis based on ribosomal DNA among 26 kinds of plant species revealed genetic similarity between the ice plant and poplar, which have salt tolerance. Overall, 35,702 protein-coding regions were identified in the genome, of which 56.05% to 82.59% were annotated and submitted to domain searches and gene ontology (GO) analyses, which found that eighteen GO terms stood out among five plant species. These terms were related to biological defense, growth, reproduction, transcription, post-transcription, and intermembrane transportation, regarded as one of the fundamental results of using the utilized ice plant genome. Conclusions The information that we characterized is useful for elucidation of the mechanism of growth promotion under salinity and reversible conversion of the photosynthetic type from C3 to Crassulacean Acid Metabolism (CAM).

Overexpressing plant ferredoxin-like protein enhances photosynthetic efficiency and carbohydrates accumulation in Phalaenopsis.

Crassulacean acid metabolism (CAM) is one of three major models of carbon dioxide assimilation pathway with better water-use efficiency and slower photosynthetic efficiency in photosynthesis. Previous studies indicated that the gene of sweet pepper plant ferredoxin-like protein (PFLP) shows high homology to the ferredoxin-1(Fd-1) family that belongs to photosynthetic type Fd and involves in photosystem I. It is speculated that overexpressing pflp in the transgenic plant may enhance photosynthetic efficiency through the electron transport chain (ETC). To reveal the function of PFLP in photosynthetic efficiency, pflp transgenic Phalaenopsis, a CAM plant, was generated to analyze photosynthetic markers. Transgenic plants exhibited 1.2-folds of electron transport rate than that of wild type (WT), and higher CO2 assimilation rates up to 1.6 and 1.5-folds samples at 4pm and 10pm respectively. Enzyme activity of phosphoenolpyruvate carboxylase (PEPC) was increased to 5.9-folds in Phase III, and NAD+-linked malic enzyme (NAD+-ME) activity increased 1.4-folds in Phase IV in transgenic plants. The photosynthesis products were analyzed between transgenic plants and WT. Soluble sugars contents such as glucose, fructose, and sucrose were found to significantly increase to 1.2, 1.8, and 1.3-folds higher in transgenic plants. The starch grains were also accumulated up to 1.4-folds in transgenic plants than that of WT. These results indicated that overexpressing pflp in transgenic plants increases carbohydrates accumulation by enhancing electron transport flow during photosynthesis. This is the first evidence for the PFLP function in CAM plants. Taken altogether, we suggest that pflp is an applicable gene for agriculture application that enhances electron transport chain efficiency during photosynthesis.

A comparison of stomatal conductance responses to blue and red light between C3 and C4 photosynthetic species in three phylogenetically-controlled experiments.

C4 photosynthesis is an adaptation that has independently evolved at least 66 times in angiosperms. C4 plants, unlike their C3 ancestral, have a carbon concentrating mechanism which suppresses photorespiration, often resulting in faster photosynthetic rates, higher yields, and enhanced water use efficiency. Moreover, the presence of C4 photosynthesis greatly alters the relation between CO2 assimilation and stomatal conductance. Previous papers have suggested that the adjustment involves a decrease in stomatal density. Here, we tested if C4 species also have differing stomatal responses to environmental cues, to accommodate the modified CO2 assimilation patterns compared to C3 species. To test this hypothesis, stomatal responses to blue and red-light were analysed in three phylogenetically linked pairs of C3 and C4 species from the Cleomaceae (Gynandropsis and Tarenaya), Flaveria, and Alloteropsis, that use either C3 or C4 photosynthesis. The results showed strongly decreased stomatal sensitivity to blue light in C4 dicots, compared to their C3 counterparts, which exhibited significant blue light responses. In contrast, in C3 and C4 subspecies of the monocot A. semialata, the blue light response was observed regardless of photosynthetic type. Further, the quantitative red-light response varied across species, but the presence or absence of a significant stomatal red-light response was not directly associated with differences in photosynthetic pathway. Interestingly, stomatal density and morphology patterns observed across the three comparisons were also not consistent with patterns commonly asserted for C3 and C4 species. The strongly diminished blue-light sensitivity of stomatal responses in C4 species across two of the comparisons suggests a common C4 feature that may have functional implications. Altogether, the strong prevalence of species-specific effects clearly emphasizes the importance of phylogenetic controls in comparisons between C3 and C4 photosynthetic pathways.

The importance of C3 and C4 grasses and CAM shrubs in the Greater Cape Floristic Region under contemporary and Last Glacial Maximum climates

Southern Africa’s Greater Cape Floristic Region (GCFR) – a hotspot of biological diversity and palaeo-anthropological evidence for modern human evolution – is a climatically complex region where broad climatic gradients influence the relative importance of plant photosynthetic types in local vegetation. This mix of photosynthetic pathways imprint an isotopic signature on palaeo-records, thereby allowing for the reconstruction of historical climate and vegetation. Here, we use vegetation plot data, coupled with the photosynthetic-type affinities of plant genera, to relate their relative cover to bioclimatic drivers and use these to create spatial distribution models of C3 and C4 grasses and CAM shrubs across the contemporary GCFR. We then use palaeodistribution modelling, drawing on downscaled climate models for the Last Glacial Maximum (LGM), to hindcast the LGM distributions of photosynthetic types for the GCFR and its then-exposed offshore areas. In addition, we construct a model to account for the effect of lower atmospheric CO2 concentration ([CO2]) during the LGM on the competitive interactions between C3 and C4 grasses and use this model to hindcast relative C4-grass cover across the glacial-era GCFR. Our palaeodistribution models suggest that, even though there were some reductions in CAM cover and shifts in C3-/C4-dominated grass layers between the LGM and Holocene, these were mostly localised, with changes at the regional scale being relatively muted. For C3/C4 grasses, distributional shifts were even more subtle once sub-ambient glacial [CO2] was accounted for. Our hindcasts of glacial, [CO2]-adjusted grass cover show agreements and inconsistencies with various sedimentary and herbivore-diet records in the palaeo-archive: both lines of evidence suggest the dominance of C3 species in the strongly winter-rainfall western GCFR; in the eastern GCFR, with muted rainfall seasonality, hindcasts of relative C3:C4 cover and δ13C signals from palaeo-records vary, with greater disparities between them. The lack of significant upheavals in the distribution of photosynthetic types suggests that the GCFR presented a relatively stable and dependable resourcescape for hunter-gatherer populations through the glacial–interglacial cycle of the LGM–Holocene transition. Our maps of relative C3-/C4-grass and CAM-shrub cover provide a reasonable surrogate for relative biomass of different photosynthetic types in local vegetation at a landscape scale and depict an independent estimate of carbon isoscapes across the GCFR – under both an interglacial (Holocene) and glacial (terminal Pleistocene) climate – against which palaeo-records can be assessed.

Soil organic carbon stock change following perennialization: a meta-analysis

Perennial crops replacing annual crops are drawing global attention because they harbor potential for sustainable biomass production and climate change mitigation through soil carbon sequestration. At present, it remains unclear how long perennial crops can sequester carbon in the soil and how soil carbon stock dynamics are influenced by climate, soil, and plant properties across the globe. This study presents a meta-analysis synthesizing 51 publications (351 observations at 77 sites) distributed over different pedo-climatic conditions to scrutinize the effect of perennialization on organic carbon accumulation in soil compared with two annual benchmark systems (i.e., monoculture and crop rotation). Results showed that perennial crops significantly increased soil organic carbon stock by 16.6% and 23.1% at 0–30 cm depth compared with monoculture and crop rotation, respectively. Shortly after establishment (< 5 years), perennial crops revealed a negative impact on soil organic carbon stock; however, long duration (> 10 years) of perennialization had a significant positive effect on soil organic carbon stock by 30% and 36.4% at 0–30 cm depth compared with monoculture and crop rotation, respectively. Compared with both annual systems, perennial crops significantly increased soil organic carbon stock regardless of their functional photosynthetic types (C3, C4, or C3-C4 intermediates) and vegetation type (woody or herbaceous). Among other factors, pH had a significant impact on soil organic carbon; however, the effect of soil textures showed no significant impact, possibly due to a lack of observations from each textural class and mixed pedoclimatic effects. Results also showed that time effect of perennialization revealed a sigmoidal increase of soil organic carbon stock until about 20 years; thereafter, the soil carbon stocks advanced towards a steady-state level. In conclusion, perennial crops increased soil organic carbon stock compared with annual systems; however, the time since conversion from annual to perennial system decisively impacted soil organic carbon stock changes.

The first released available genome of the common ice plant ( Mesembryanthemum crystallinum L.) extended the research region on salt tolerance, C 3-CAM photosynthetic conversion, and halophilism.

The common ice plant ( Mesembryanthemum crystallinum L.) is an annual herb belonging to the genus Mesembryanthemum of the family Aizoaceae, native to Southern Africa. We performed shotgun genome paired-end sequencing using the Illumina platform to determine the genome sequence of the ice plants. We assembled the whole genome sequences using the genome assembler "ALGA" and "Redundans", then released them as available genomic information. Finally, we mainly estimated the potential genomic function by the homology search method. A draft genome was generated with a total length of 286 Mb corresponding to 79.2% of the estimated genome size (361 Mb), consisting of 49,782 contigs. It encompassed 93.49% of the genes of terrestrial higher plants, 99.5% of the ice plant transcriptome, and 100% of known DNA sequences. In addition, 110.9 Mb (38.8%) of repetitive sequences and untranslated regions, 971 tRNA, and 100 miRNA loci were identified, and their effects on stress tolerance and photosynthesis were investigated. Molecular phylogenetic analysis based on ribosomal DNA among 26 kinds of plant species revealed genetic similarity between the ice plant and poplar, which have salt tolerance. Overall, 35,702 protein-coding regions were identified in the genome, of which 56.05% to 82.59% were annotated and submitted to domain searches and gene ontology (GO) analyses, which found that eighteen GO terms stood out among five plant species. These terms were related to biological defense, growth, reproduction, transcription, post-transcription, and intermembrane transportation, regarded as one of the fundamental results of using the utilized ice plant genome. The information that we characterized is useful for elucidation of the mechanism of growth promotion under salinity and reversible conversion of the photosynthetic type from C3 to Crassulacean Acid Metabolism (CAM).

Alloteropsis semialata as a study system for C4 evolution in grasses.

Numerous groups of plants have adapted to CO2 limitations by independently evolving C4 photosynthesis. This trait relies on concerted changes in anatomy and biochemistry to concentrate CO2 within the leaf and thereby boost productivity in tropical conditions. The ecological and economic importance of C4 photosynthesis has motivated intense research, often relying on comparisons between distantly related C4 and non-C4 plants. The photosynthetic type is fixed in most species, with the notable exception of the grass Alloteropsis semialata. This species includes populations exhibiting the ancestral C3 state in southern Africa, intermediate populations in the Zambezian region and C4 populations spread around the palaeotropics. We compile here the knowledge on the distribution and evolutionary history of the Alloteropsis genus as a whole and discuss how this has furthered our understanding of C4 evolution. We then present a chromosome-level reference genome for a C3 individual and compare the genomic architecture with that of a C4 accession of A. semialata. Alloteropsis semialata is one of the best systems in which to investigate the evolution of C4 photosynthesis because the genetic and phenotypic variation provides a fertile ground for comparative and population-level studies. Preliminary comparative genomic investigations show that the C3 and C4 genomes are highly syntenic and have undergone a modest amount of gene duplication and translocation since the different photosynthetic groups diverged. The background knowledge and publicly available genomic resources make A. semialata a great model for further comparative analyses of photosynthetic diversification.

A Machine Learning Framework Identifies Plastid-Encoded Proteins Harboring C3 and C4 Distinguishing Sequence Information.

C4 photosynthesis is known to have at least 61 independent origins across plant lineages making it one of the most notable examples of convergent evolution. Of the >60 independent origins, a predicted 22-24 origins, encompassing greater than 50% of all known C4 species, exist within the Panicoideae, Arundinoideae, Chloridoideae, Micrairoideae, Aristidoideae, and Danthonioideae (PACMAD) clade of the Poaceae family. This clade is therefore primed with species ideal for the study of genomic changes associated with the acquisition of the C4 photosynthetic trait. In this study, we take advantage of the growing availability of sequenced plastid genomes and employ a machine learning (ML) approach to screen for plastid genes harboring C3 and C4 distinguishing information in PACMAD species. We demonstrate that certain plastid-encoded protein sequences possess distinguishing and informative sequence information that allows them to train accurate ML C3/C4 classification models. Our RbcL-trained model, for example, informs a C3/C4 classifier with greater than 99% accuracy. Accurate prediction of photosynthetic type from individual sequences suggests biologically relevant, and potentially differing roles of these sequence products in C3 versus C4 metabolism. With this ML framework, we have identified several key sequences and sites that are most predictive of C3/C4 status, including RbcL, subunits of the NAD(P)H dehydrogenase complex, and specific residues within, further highlighting their potential significance in the evolution and/or maintenance of C4 photosynthetic machinery. This general approach can be applied to uncover intricate associations between other similar genotype-phenotype relationships.

The first released available genome of the common ice plant (Mesembryanthemum crystallinum L.) extended the research region on salt tolerance, C3-CAM photosynthetic conversion, and halophism

Background: The common ice plant (Mesembryanthemum crystallinum L.) is an annual herb belonging to the genus Mesembryanthemum of the family Aizoaceae, native to Southern Africa. Methods: We performed shotgun genome paired-end sequencing using the Illumina platform to determine the genome sequence of the ice plants. We assembled the whole genome sequences using the genome assembler “ALGA” and “Redundans”, then released them as available genomic information. Finally, we mainly estimated the potential genomic function by the homology search method. Results: A draft genome was generated with a total length of 286 Mb corresponding to 79.2% of the estimated genome size (361 Mb), consisting of 49,782 contigs. It encompassed 93.49% of the genes of terrestrial higher plants, 99.5% of the ice plant transcriptome, and 100% of known DNA sequences. In addition, 110.9 Mb (38.8%) of repetitive sequences and untranslated regions, 971 tRNA, and 100 miRNA loci were identified, and their effects on stress tolerance and photosynthesis were investigated. Molecular phylogenetic analysis based on ribosomal DNA among 26 kinds of plant species revealed genetic similarity between the ice plant and poplar, which have salt tolerance. Overall, 35,702 protein-coding regions were identified in the genome, of which 56.05% to 82.59% were annotated and submitted to domain searches and gene ontology (GO) analyses, which found that eighteen GO terms stood out among five plant species. These terms were related to biological defense, growth, reproduction, transcription, post-transcription, and intermembrane transportation, regarded as one of the fundamental results of using the utilized ice plant genome. Conclusions: The information that we characterized is useful for elucidation of the mechanism of growth promotion under salinity and reversible conversion of the photosynthetic type from C3 to Crassulacean Acid Metabolism (CAM).

The contemporary distribution of grasses in Australia: A process of immigration, dispersal and shifting dominance

AbstractAimLittle is known about the distribution of grasses throughout Australia. Using endemism as a basis for understanding biogeographical distributions, we hypothesised that contemporary species richness would be the result of environmental factors and dynamic ecological interactions spanning more than 25 Ma.LocationAustralia.TaxonGrasses.MethodsWe mapped the distribution of all Australian grass species and modelled climatic and landscape correlates according to photosynthetic type (C3 or C4), endemism, age in Australia, phylogenetic lineage and traits linked to dominance, using height as a proxy. Age classes comprised ‘Ancient’ (Gondwanan), and three others related to migration during the Sunda‐Sahul Interchange (SSI): Early, Mid or Recent. In some analyses, ‘Ancient’, ‘Early SSI’ and ‘Mid SSI’ were combined into ‘Pre‐Recent SSI’.ResultsOverall, species richness of C4 grasses increased with warmer mean annual temperatures, while richness of C3 grasses was higher in cooler areas. Recent SSI species had strong associations with summer rains and were dominant in the continent's northeast, with Pre‐Recent SSI species concentrated in the southeast, a pattern largely reflecting photosynthetic type (C4 and C3 respectively). Endemic and shared species distribution patterns support a migration sequence in which most C3 Pooideae and Panicoideae genera arrived in Australia before the Pliocene aridifications, followed by C4 Chloridoideae as aridification increased, with C4 Andropogoneae immigrating most recently across Lake Carpentaria's open habitats in the later Pleistocene. Recent SSI shared species were significantly taller than Pre‐Recent SSI endemic grasses.Main ConclusionsThe few grasses present in Australia before the Pliocene grew in cooler areas. The influx of taller Recent SSI grasses contributed to dramatic environmental changes—including creation of the northern savannas—with repercussions for resident taxa. Contemporary methods of fuel management could be promoting invasion by grass, thereby jeopardising the conditions suited to ancient taxa and threatening the region's evolutionary history.

Relationships of the competitor, stress tolerator, ruderal functional strategies of grass species with lifespan, photosynthetic type, naturalization and climate.

Grass species (family Poaceae) are globally distributed, adapted to a wide range of climates and express a diversity of functional strategies. We explored the functional strategies of grass species using the competitor, stress tolerator, ruderal (CSR) system and asked how a species' strategy relates to its functional traits, climatic distribution and propensity to become naturalized outside its native range. We used a global set of trait data for grass species to classify functional strategies according to the CSR system based on leaf traits. Differences in strategies in relation to lifespan (annual or perennial), photosynthetic type (C3 or C4), or naturalisation (native or introduced) were investigated. In addition, correlations with traits not included in the CSR classification were analyzed, and a model was fitted to predict a species' average mean annual temperature and annual precipitation across its range as a function of CSR scores. Values for competitiveness were higher in C4 species than in C3 species, values for stress tolerance were higher in perennials than in annuals, and introduced species had more pronounced competitive-ruderal strategies than native species. Relationships between the CSR classification, based on leaf traits, and other functional traits were analyzed. Competitiveness was positively correlated with height, while ruderality was correlated with specific root length, indicating that both above- and belowground traits underlying leaf and root economics contribute to realized CSR strategies. Further, relationships between climate and CSR classification showed that species with competitive strategies were more common in warm climates and at high precipitation, whereas species with stress tolerance strategies were more common in cold climates and at low precipitation. The findings presented here demonstrate that CSR classification of functional strategies based on leaf traits matches expectations for the adaptations of grass species that underlie lifespan, photosynthetic type, naturalization and climate.

The first released available genome of the common ice plant (Mesembryanthemum crystallinum L.) extended the research region on salt tolerance, C3-CAM photosynthetic conversion, and halophism

Background: The common ice plant (Mesembryanthemum crystallinum L.) is an annual herb belonging to the genus Mesembryanthemum of the family Aizoaceae, native to Southern Africa. Methods: We performed shotgun genome paired-end sequencing using the Illumina platform to determine the genome sequence of the ice plants. We assembled the whole genome sequences using the genome assembler “ALGA” and “Redundans”, then released them as available genomic information. Finally, we mainly estimated the potential genomic function by the homology search method. Results: A draft genome was generated with a total length of 286 Mb corresponding to 79.2% of the estimated genome size (361 Mb), consisting of 49,782 contigs. It encompassed 93.49% of the genes of terrestrial higher plants, 99.5% of the ice plant transcriptome, and 100% of known DNA sequences. In addition, 110.9 Mb (38.8%) of repetitive sequences and untranslated regions, 971 tRNA, and 100 miRNA loci were identified, and their effects on stress tolerance and photosynthesis were investigated. Molecular phylogenetic analysis based on ribosomal DNA among 26 kinds of plant species revealed genetic similarity between the ice plant and poplar, which have salt tolerance. Overall, 35,702 protein-coding regions were identified in the genome, of which 56.05% to 82.59% were annotated and submitted to domain searches and gene ontology (GO) analyses, which found that eighteen GO terms stood out among five plant species. These terms were related to biological defense, growth, reproduction, transcription, post-transcription, and intermembrane transportation, regarded as one of the fundamental results of using the utilized ice plant genome. Conclusions: The information that we characterized is useful for elucidation of the mechanism of growth promotion under salinity and reversible conversion of the photosynthetic type from C3 to Crassulacean Acid Metabolism (CAM).

The evolution and diurnal expression patterns of photosynthetic pathway genes of the invasive alien weed, Mikania micrantha

Mikania micrantha is a fast-growing global invasive weed species that causes severe damage to natural ecosystems and very large economic losses of forest and crop production. It has advantages in photosynthesis, including a similar net photosynthetic rate as C4 plants and a higher carbon fixation capacity. We used a combination of genomics and transcriptomics approaches to study the evolutionary mechanisms and circadian expression patterns of M. micrantha. In M. micrantha, 16 positive selection genes focused on photoreaction and utilization of photoassimilates. In different tissues, 98.1% of the genes associated with photoresponse had high expression in stems, and more than half of the genes of the C4 cycle had higher expression in stems than in leaves. In stomatal opening and closing, 2 genes of carbonic anhydrase (CAs) had higher expression at 18:00 than at 8:00, and the slow anion channel 1 (SLAC1) and high-leaf-temperature 1 kinase (HT1) genes were expressed at low levels at 18:00. In addition, genes associated with photosynthesis had higher expression levels at 7:00 and 17:00. We hypothesized that M. micrantha may undergo photosynthesis in the stem and flower organs and that some stomata of the leaves were opening at night by CO2 signals. In addition, its evolution may attenuate photoinhibition at high light intensities, and enhance more efficient of photosynthesis during low light intensity. And the tissue-specific photosynthetic types and different diurnal pattern of photosynthetic-related genes may contribute to its rapid colonization of new habitats of M. micrantha.

15N Natural Abundance of C3 and C4 Herbaceous Plants and Its Response to Climatic Factors along an Agro-Pastoral Zone of Northern China.

The nitrogen isotope composition of plants (δ15N) can comprehensively reflect information on climate change and ecosystems' nitrogen cycle. By collecting common herbs and soil samples along the 400 mm isoline of mean annual precipitation (MAP) in the agro-pastoral zone of North China (APZNC) and measuring their δ15N values, the statistical characteristics of foliar δ15N of herbs and the responses of foliar δ15N to the MAP and mean annual temperature (MAT) were analyzed. The results showed that: (1) the δ15N values of all herbs investigated varied from -5.5% to 15.25%. Among them, the δ15N value range of C3 herbs (-5.5~15.00%) was wider than that of C4 herbs (-2.17~15.25%), but the average value (3.27%) of C3 herbs was significantly lower than that of C4 herbaceous plants (5.55%). This difference provides an important method for identifying plants of different photosynthetic types by nitrogen isotope technology. (2) Along the transect from northeast to southwest, the δ15N of both C3 and C4 herbs decreased with the increase in the MAP, but not significantly for C3 herbs. The inverse relationship between the nitrogen isotopic signatures of herbs and MAP is consistent with previous studies. However, the MAP in the APZNC is found to only explain a small amount of the observed variance in the δ15N herbs (C3 herbs: 10.40%; C4 herbs: 25.03%). (3) A strong negative relationship was found between δ15N of herbs and MAT across the transect (C3 herbs: -0.368%/°C; C4 herbs: -0.381%/°C), which was contrary to the global pattern and some regional patterns. There was no significant difference in the δ15N responses of two different photosynthetic herbs to temperature, but the effect of temperature on the variances of δ15N of C3 and C4 herbs was significantly greater than that of precipitation. This suggests that temperature is a key factor affecting foliar δ15N of herbs in this transect. The above findings may be of value to global change researchers studying the processes of the nitrogen cycle and gaining an insight into climate dynamics of the past.

C4 photosynthesis in Paulownia? A case of inaccurate citations

Societal Impact StatementC4 photosynthesis is an ultra‐efficient mode of photosynthesis found in some of our most productive crop species yet is notably rare in trees. Given C4 photosynthesis is associated with high yield in herbaceous species, especially under hot and dry conditions, C4 trees may seem an attractive prospect for biomass production and carbon sequestration in a rapidly changing climate. This may explain why some in the literature have optimistically linked C4 photosynthesis with the exceptionally fast‐growing tree Paulownia. However, this claim is lacking in evidence and represents an example of poor citation practices leading to the spread of misinformation.Summary The rapid growth of trees in genus Paulownia (Paulowniaceae) has been attributed in the literature to their use of C4 photosynthesis, a complex trait that confers increased photosynthetic efficiency under certain environmental conditions. After careful examination of citations used to support the idea that Paulownia species use C4 photosynthesis, we find that there is no data underpinning this claim. Despite this, many investment schemes utilise information about the physiology of Paulownia, including photosynthetic type, to legitimise the use of Paulownia trees for financial investment and carbon offsetting. This study uses leaf physiology, anatomy and stable isotope data to determine whether or not three species in Paulownia (Paulownia tomentosa, Paulownia fortunei and Paulownia kawakamii) use C4 photosynthesis. These data are compared with existing data for C3 and C4 woody species in the literature. We show that the leaf physiology, anatomy and stable isotope phenotypes of the three Paulownia trees considered in the study are not consistent with those of C4 plants. Our findings highlight how inaccurate citation of scientific findings can contribute to the spread of misinformation beyond the scientific community, as some of those promoting investments in Paulownia plantations reference the photosynthetic superiority of Paulownia as a means to legitimise its use in carbon offsetting.

Photosynthetic nitrogen- and water-use efficiencies in C3 and C4 subtype grasses grown under two nitrogen supply levels

ABSTRACT C4 plants have higher photosynthetic nitrogen- and water-use efficiencies (PNUE and PWUE) than C3 plants. Some studies on C4 grasses have reported that NADP-malic enzyme (NADP-ME) grasses have higher PNUE than NAD-malic enzyme (NAD-ME) grasses, whereas other studies have suggested that this difference is caused by the traits of phylogenetic lineage. Further extensive studies are required to properly understand the PNUE and PWUE of C4 grasses. We investigated the PNUE and PWUE for 22 grass species of four photosynthetic types [4 NADP-ME, 6 NAD-ME, 6 phosphoenolpyruvate carboxykinase (PCK), and 6 C3 species] grown under two N supply levels. Under the same N level, the three C4 types showed higher photosynthetic rate and lower stomatal conductance and transpiration rate than the C3 type, but these parameters were similar in the C4 types. Although there were no significant differences in leaf mass per area and N content per leaf area among the four types, these traits tended to be somewhat lower in the NADP-ME type than in other types. As a result, PNUE was highest in the NADP-ME type, lowest in the C3 type, and intermediate in the NAD-ME and PCK types. PWUE was much higher in the C4 types than in the C3 type but similar in the C4 types. The N supply level did not greatly affect the trends of PNUE and PWUE. This study suggests that the NAD-ME and PCK grasses have the N-related physiological traits similar to each other but differing from the NADP-ME grasses.

Insights into Regulation of C2 and C4 Photosynthesis in Amaranthaceae/Chenopodiaceae Using RNA-Seq.

Amaranthaceae (incl. Chenopodiaceae) shows an immense diversity of C4 syndromes. More than 15 independent origins of C4 photosynthesis, and the largest number of C4 species in eudicots signify the importance of this angiosperm lineage in C4 evolution. Here, we conduct RNA-Seq followed by comparative transcriptome analysis of three species from Camphorosmeae representing related clades with different photosynthetic types: Threlkeldia diffusa (C3), Sedobassia sedoides (C2), and Bassia prostrata (C4). Results show that B. prostrata belongs to the NADP-ME type and core genes encoding for C4 cycle are significantly upregulated when compared with Sed. sedoides and T. diffusa. Sedobassia sedoides and B. prostrata share a number of upregulated C4-related genes; however, two C4 transporters (DIT and TPT) are found significantly upregulated only in Sed. sedoides. Combined analysis of transcription factors (TFs) of the closely related lineages (Camphorosmeae and Salsoleae) revealed that no C3-specific TFs are higher in C2 species compared with C4 species; instead, the C2 species show their own set of upregulated TFs. Taken together, our study indicates that the hypothesis of the C2 photosynthesis as a proxy towards C4 photosynthesis is questionable in Sed. sedoides and more in favour of an independent evolutionary stable state.