CO 2-induced chloroplast movement was reported in the monograph by Gustav Senn in 1908: unilateral CO 2 supply to the one cell-layered moss leaves induced the positively CO 2-tactic periclinal arrangement of chloroplasts. However, from the modern criteria, several experimental settings are unacceptable. Here, using a model moss plant Physcomitrium patens, we examined basic features of chloroplast CO 2-tactic relocation with a modernized experimental system. The CO 2 relocation was light-dependent and especially the CO 2 relocation in red light was substantially dependent on photosynthetic activity. Between the cytoskeletons responsible for chloroplast movement of P. patens, the microfilament mainly worked for CO 2 relocation, but the microtubule-based movement was insensitive to CO 2. The CO 2 relocation was induced not only by air with and without CO 2 but also by the more realistic difference in CO 2 concentration between the two sides. In the leaves placed on the surface of a gel sheet, chloroplasts avoided the gel side and positioned in the air facing surface. This was also shown to be photosynthesis dependent. Based on these observations, we propose a working hypothesis that the threshold light intensity between the light-accumulation and -avoidance responses of the photorelocation would be increased by CO 2, resulting in the CO 2-tactic relocation of chloroplast.
The combined study of C and O isotopes in plant organic matter has emerged as a powerful tool for understanding plant functional responses to environmental change. The approach relies on established relationships between leaf gas exchange and isotopic fractionation to derive a series of model scenarios that can be used to infer changes in photosynthetic assimilation and stomatal conductance driven by changes in environmental parameters (CO2, water availability, air humid-ity, temperature, nutrients). We review the mechanistic basis for a conceptual model, in light of recently published research, and discuss where isotopic observations don’t match our current understanding of plant physiological response to environment. We demonstrate that 1) the mod-el was applied successfully in many, but not all studies, 2), while originally conceived for leaf isotopes, the model has been applied extensively to tree ring isotopes in the context of tree physiology and dendrochronology. Where isotopic observations deviate from physiologically plau-sible conclusions, this mismatch between gas-exchange and isotope response provides valuable insights on underlying physiological processes. Overall, we found that isotope responses can be grouped into situations of increasing resource limitation versus higher resource availability. The dual isotope model helps to interpret plant responses to a multitude of environmental factors.
Stomata are the key nodes linking photosynthesis and transpiration. By regulating the opening degree of stomata, plants successively achieve the balance between water loss and carbon dioxide acquisition. The dynamic behavior of stomata is an important cornerstone of plant adaptability. Though there have been miscellaneous experimental results on stomata and their constituent cells, the guard cells and the subsidiary cells, current theory of stomata regulation is far from clear and unified. In this work, we develop an integrated model to describe the stomatal dynamics of seed plants based on existing experimental results. The model includes three parts: 1) a passive mechanical model of the stomatal aperture as a bivariate function of the guard-cell and the subsidiary-cell turgor pressures; 2) an active regulation model with a targeted ion-content in guard cells as a function of their water potential; and 3) a dynamical model for the movement of potassium ions and water content. Our model has been used to reproduce different experimental phenomena semi and stomatal responses to environment conditions.
Conservative flowering behaviors, such as flowering during long days in summer or late flowering at a high leaf number, are often proposed to protect against variable winter and spring temperatures which lead to frost damage if premature flowering occurs. Yet, due the many factors in natural environments relative to the number of individuals compared, assessing which climate characteristics drive these flowering traits has been difficult. We applied a multidisciplinary approach to ten winter-annual Arabidopsis thaliana populations originating along a wide climactic gradient in Norway. We used a variable reduction strategy to assess which of 100 climate descriptors from their home sites correlated most to their behaviors when grown in common garden and assessed sequence variation of 19 known environmental-response flowering genes. Photoperiod sensitivity inversely correlated with interannual variation in timing of growing season onset (start of favorable spring temperatures). Time to flowering appeared driven by growing season length, curtailed by cold fall temperatures. The distribution of FLM, TFL2, and HOS1 haplotypes, genes involved in ambient temperature response, correlated with growing-season climate. We show that long-day sensitivity and late flowering may be driven not by risk of spring frosts, but by growing season temperature and length perhaps to opportunistically maximize growth.
β-Glucosidase is validated as an elicitor for early immune responses in plants and it was detected in the salivary glands of Frankliniella occidentalis in previous research. Seven differentially expressed genes encoding β-Glucosidase were obtained by comparing the transcriptomes of F. occidentalis adults grown under two different CO 2 concentrations (800 ppm vs. 400 ppm), which might be associated with the differences in the interaction between F. occidentalis adults and its host plant, Phaseolus vulgaris under different CO 2 levels. To verify this speculation, changes in defense responses based on the production and elimination of reactive oxygen species (ROS) in P. vulgaris leaves treated with three levels of β-Glucosidase activity under ambient CO 2 (aCO 2) and elevated CO 2 (eCO 2) were measured in this study. The results showed that both leaves infested with thrips and those sprayed with the pure β-Glucosidase solution showed significant increases in ROS levels under aCO 2 and eCO 2, and the activities of antioxidant enzymes including superoxide dismutase (SOD), peroxidase (POD), and catalase (CAT) were increased correspondingly, while in leaves infested with FoβGlu-1-silenced thrips, the ROS levels and activities of these enzymes did not change significantly during the first 12 hours of injury regardless of CO 2 level. Besides, significantly higher levels of ROS and lower activities of SOD, POD and CAT in injured leaves under eCO 2 compared to aCO 2 were noticed, which would negatively affect P. vulgaris leaves and facilitate thrips damage.
As sessile organisms, plants are constantly challenged by a dynamic growing environment. This includes fluctuations in temperature, water availability, light levels, and atmospheric conditions. In concert with changes in abiotic conditions, plants experience changes in biotic stress pressures, including plant pathogens, viruses, and herbivores. Human-induced increases in atmospheric carbon dioxide (CO 2) levels have led to alterations in plant growth environments that challenge their productivity and nutritional quality. Additionally, it is predicted that climate change will alter the prevalence and virulence of plant pathogens, further challenging plant productivity. A knowledge gap exists in the complex interplay between plant responses to biotic and abiotic stress conditions. Closing this gap is crucial for developing climate resilient crops in the future. Here, we review the physiological responses of plants to elevated CO 2, temperature, tropospheric ozone (O 3), and drought conditions, as well as the interaction of these abiotic stress factors with plant pathogen pressure. Additionally, we describe the crosstalk and trade-offs involved in plant responses to both abiotic and biotic stress, and outline targets for future work to develop a more sustainable future food supply in light of future climate change.
The linkage of stomatal behavior with photosynthetic processes is critical to understanding water and carbon cycles under global change. The slope ( g1) of stomatal conductance ( gs) versus CO 2 assimilation ( Anet) serves as a proxy of the marginal water cost of carbon acquisition and the trade-off between carbon gain and water loss. Here we use g1 to assess species differences in the response of stomatal behavior to experimental climate change manipulations, asking whether generalizable patterns exist across species and climate contexts. A total of 17,727 Anet- gs measurements made in a long-term open-air experiment under ambient and +3.3°C warming, and ambient and ~40% summer rainfall reduction provided > 2,700 estimates of g1 across 21 boreal and temperate tree species. All species became more conservative in their water use (lower g1) in warming and/or reduced rainfall treatments because of lower soil moisture. In contrast to these phenotypic responses, species from warmer and drier habitats tended to have slightly higher g1 and to be the least sensitive to the decrease in soil water. Overall, both warming and rainfall reduction consistently made stomatal behavior more conservative in terms of water loss per unit carbon gain across 21 species and a decade of experimental observation.
Local adaptation is a major driver of biological diversity, and related species may develop analogous (parallel evolution) or alternative (divergent evolution) solutions to similar ecological challenges. We expect these adaptive solutions between closely related organisms would culminate in both phenotypic and genotypic signals. In this study, we employ a reciprocal transplant, glasshouse experiment with two Eucalyptus species ( E. grandis and E. tereticornis) with large, overlapping distributions grown under contrasting ‘local’ temperature conditions (tropic and temperate) to investigate the independent contribution of adaptation, plasticity, and their interaction at molecular, physiological and morphological levels. We find key traits differ in their response. The link between gene expression and traits markedly differed between species. Divergent evolution was the dominant pattern driving adaptation as unique gene responses (91% of all significant genes) was the greatest factor driving differentiation; but overlapping gene (homologous) responses were dependent on the determining factor (plastic, adaptive, or genotype by environment interaction). 98% of the plastic homologs were similarly regulated, while 50% of the adaptive homologs and 100% of the interaction homologs were antagonistically regulated. Therefore, parallel evolution for the adaptive effect in homologous genes was greater than expected but not in favour of divergent evolution. Further, heat shock proteins for E. grandis were almost entirely driven by adaptive responses, while plasticity drove the response in E. tereticornis. These results suggest divergent molecular evolutionary solutions dominated the adaptive mechanisms among species, even in similar ecological circumstances. Thus, trees with overlapping distributions are unlikely to equally persist in the future, suggesting that management of future forests to changing temperature conditions must be species specific.
Group VII Ethylene Response Factors (ERF-VII) are plant-specific transcription factors (TFs) known for their role in the activation of hypoxia-responsive genes under low oxygen stress but also in plant endogenous hypoxic niches. However, their function in the microaerophilic nitrogen-fixing nodules of legumes has not yet been investigated. We investigated regulation and the function of the two Medicago truncatula ERF-VII TFs ( MtERF74 and MtERF75) in roots and nodules, MtERF74 and MtERF75 in response to hypoxia stress and during the nodulation process using an RNA interference strategy and targeted proteolysis of MtERF75. Knockdown of MtERF74 and MtERF75 partially blocked the induction of hypoxia-responsive genes in roots exposed to hypoxia stress. In addition, a significant reduction in nodulation capacity and nitrogen fixation activity was observed in mature nodules of double knockdown transgenic roots. Overall, the results indicate that MtERF74 and MtERF75 are involved in the induction of MtNR1 and Pgb1.1 expression for efficient Phytogb-NO respiration in the nodule.
Plant metabolomics has been used widely in plant physiology, in particular to analyse metabolic responses to environmental parameters. Derivatization (via trimethylsilylation-methoximation) followed by GC-MS metabolic profiling is a major technique to quantify low molecular weight, common metabolites of primary carbon, sulphur and nitrogen metabolism. There are now excellent opportunities for new generation analyses, using high resolution, exact mass GC-MS spectrometers that are progressively becoming relatively cheap. However, exact mass GC-MS analyses for routine metabolic profiling are not common, there is no dedicated available database. Also, exact mass GC-MS is usually dedicated to structural resolution of targeted secondary metabolites. Here, we present a curated database for exact mass metabolic profiling (made of 336 analytes, 1,064 characteristic exact mass fragments) focused on molecules of primary metabolism. We show advantages of exact mass analyses, in particular to resolve isotopic patterns, localise S-containing metabolites, and avoid identification errors when analytes have common nominal mass peaks in their spectrum. We provide a practical example using leaves of different Arabidopsis ecotypes and show how exact mass GC-MS analysis can be applied to plant samples and identify metabolic profiles.
Triose-phosphate utilization (TPU) limits the maximum rate at which plants can photosynthesize. However, TPU is almost never found to be limiting photosynthesis under ambient conditions for plants. This, along with previous results showing adaptability of TPU at low temperature, suggest that TPU capacity is regulated to be just above the photosynthetic rate achievable under the prevailing conditions. A set of experiments were performed to study the adaptability of TPU capacity when plants are acclimated to elevated CO 2 concentrations. Plants held at 1500 ppm CO 2 were initially TPU limited. After 30 hours they no longer exhibited TPU limitations but they did not elevate their TPU capacity. Instead, the maximum rates of carboxylation and electron transport declined. A timecourse of regulatory responses was established. A step increase of CO 2 first caused PSI to be oxidized but after 40 s both PSI and PSII had excess electrons as a result of acceptor-side limitations. Electron flow to PSI slowed and the proton motive force increased. Eventually, non-photochemical quenching reduced electron flow sufficiently to balance the TPU limitation. Over several minutes rubisco deactivated contributing to regulation of metabolism to overcome the TPU limitation.
N-terminal cysteine oxidases (NCOs) use molecular oxygen to oxidize the amino-terminal cysteine of specific proteins, thereby initiating the proteolytic N-degron pathway. To expand the characterization of the plant family of NCOs (PCOs), we performed a phylogenetic analysis across different taxa in terms of sequence similarity and transcriptional regulation. Based on this survey, we propose a distinction of PCOs into two main groups. A-type PCOs are conserved across all plant species and are generally unaffected at the mRNA level by oxygen availability. Instead, B-type PCOs differentiated in spermatophytes to acquire transcriptional regulation in response to hypoxia. The inactivation of two A-type PCOs in Arabidopsis thaliana, PCO4 and PCO5, is sufficient to activate the anaerobic response in young seedlings, whereas the additional removal of B-type PCOs leads to a stronger induction of anaerobic genes and impairs plant growth and development. Our results show that both PCO types are required to regulate the anaerobic response in angiosperms. Therefore, while it is possible to distinguish two clades within the PCO family, we conclude that they all contribute to restrain the anaerobic transcriptional program in normoxic conditions and together generate a molecular switch to toggle the hypoxic response.
Microbe associated molecular pattern (MAMP) triggered immunity research has traditionally centred around signal transduction pathways originating from activated membrane localised pattern recognition receptors (PRRs), culminating in nuclear transcription and post translational modifications. More recently, chloroplasts have emerged as key immune signalling hubs. Chloroplasts play a central role in integrating environmental signals. Notably MAMP recognition induces chloroplastic ROS (cROS) which is suppressed by pathogens effectors, which also modify the balance of defence hormone precursors, jasmonic acid (JA), salicylic acid (SA) and abscisic acid (ABA), whose precursors are chloroplast synthesised. This study focuses on how well characterised PRRs and co-receptors modulate chloroplast physiology, examining whether diverse signalling pathways converge to similarly modulate chloroplast function. Pre-treatment of receptor mutant plants with MAMP and D(Damage)AMP peptides usually protect against effector modulation of chlorophyll fluorescence and prevent Pseudomonas syringae effector mediated quenching of cROS and suppression of Fv/Fm. The MAMP-triggered immunity (MTI) co-receptor double mutant, bak1-5/bkk1-1, exhibits a remarkable decrease in Fv/Fm compared to control plants during infection, underlining the importance of MTI mediated signalling in chloroplast immunity. Further probing the role of the chloroplast in immunity we unexpectedly found that high light uncouples plant immune signalling.
Breeding drought stress tolerance is an integral part of our current and future goals of sustainable agricultural production. In the present study, we examined the natural variation of HvP5cs1 and demonstrated the utility of a wild barley allele for drought stress adaptation in cultivated barley. Sequencing the 5-end regulatory region among 49 barley accessions identified a genetically distinct allele of HvP5cs1 promoter from a wild barley ISR42-8. Allele mining of HvP5cs1 indicated quantitative variation in proline accumulation which was associated with promoter polymorphisms across the cluster of abscisic acid-responsive elements (ABRE), ABRE-related coupling elements, and MYB binding motifs. A near-isogenic line (NIL-143) harboring the HvP5cs1 allele from the highest proline accumulating wild barley ISR42-8 was developed in cultivated barley Scarlett through marker-assisted backcrossing (BC6). NIL-143 preserved the genetic competence of ISR42-8 to accumulate proline in higher concentrations under drought conditions at seedling and reproductive stages. Under drought stress, NIL-143 maintained superior membrane integrity, reduced pigment damage, and sustained photosynthetic health compared to Scarlett. NIL-143 presented a remarkable improvement in drought stress recovery than Scarlett. Further, the introgression line exhibited improved yield attributes, especially superior grain weight compared to Scarlett under field drought conditions. In conclusion, the present data uncover the genetic regulation of HvP5cs1 mediated proline accumulation and elucidate its role in drought stress adaptation and yield stability in barley.
Calcium (Ca 2+) is an important second messenger in plants. The activation of Ca 2+ signaling cascades is critical in the activation of adaptive processes in response to perceived environmental stimuli, including biotic stresses. The colonization of roots by the plant growth promoting endophyte Serendipita indica involves the increase of cytosolic Ca 2+ levels in Arabidopsis thaliana. In this study, we investigated transcriptional changes in Arabidopsis roots during symbiosis with S. indica. RNA-seq profiling disclosed the significant induction of CALCINEURIN B-LIKE 7 ( CBL7) during early- and later phases of the interaction. Consistent with the transcriptomics analysis, reverse genetic evidence and yeast two-hybrid studies highlighted the functional relevance of CBL7 and tested the involvement of a CBL7-CBL-INTERACTING PROTEIN KINASE 13 (CIPK13) signaling pathway in the establishment of the mutualistic relationship that promotes plant growth. The loss-of-function of CBL7 abolished the growth promoting effect of S. indica and affected the colonization of the root by the fungus. The subsequent transcriptomics analysis of cbl7 revealed the involvement of this Ca 2+ sensor in activating plant defense responses. Furthermore, we report on the contribution of CBL7 to potassium transport in Arabidopsis. Triggered by the differential expression of a small number of K + channels/transporter genes, we analyzed K + contents in wild-type and cbl7 plants and observed a significant accumulation of K + in root of cbl7 plants, while shoot tissues demonstrated K + depletion. Taken together, our work associates CBL7 with an important role in the mutual interaction between Arabidopsis and S. indica and links the CBL7 Ca 2+ receptor protein to K + transport.
Plant vacuoles serve as the primary intracellular compartments for phosphorus (P) storage and play a central role to maintain P homeostasis. The Oryza sativa (rice) genome contains three genes that encode SPX (SYG1/PHO81/XPR1)-MFS (Major Facility Superfamily) proteins (OsSPX-MFS1, 2, 3). OsSPX-MFS1 and OsSPX-MFS3 were shown previously to have vacuolar phosphate (Pi) transporter activities, but the physiological role of the three transporters under varying P conditions and under field grown conditions for a crop plant is not known. To address this knowledge gap, we generated single, double, and triple mutants (7 mutants with at least two lines of each) for the three rice Os SPX-MFS genes. All the mutants except osspx-mfs2 display lower vacuolar Pi concentrations and all Os SPX-MFSs overexpression plant lines display higher Pi accumulation, demonstrating that all three OsSPX-MFSs are vacuolar Pi influx transporters. OsSPX-MFS3 plays the dominant role based on the phenotypes of three single mutants in terms of growth, vacuolar and tissue Pi concentrations. OsSPX-MFS2 is the weakest and only functions as vacuole Pi sequestration under osspx-mfs1/3 background. The vacuolar Pi sequestration was severely impaired in osspx-mfs1/ 3 and osspx-mfs1/2/ 3, which led to Pi toxicity and subsequently increased Pi allocation to aerial organs. High Pi in the panicle result in necrotic symptoms on husks and impaired panicle and grain development in osspx-mfs1/ 3 and osspx-mfs1/2/ 3 mutant lines. The mutation in the weak vacuolar Pi transporter OsSPX-MFS2 resulted more stable yield compared to the wildtype under low P field conditions. The results suggest that alteration of vacuolar Pi sequestration may be a novel effective strategy to improve rice (crop) tolerance to low phosphorus field conditions and maintain yield.
Recent progress has shown that vacuolar Pi transporters (VPTs) are important for cellular Pi homeostasis against external Pi variations in Arabidopsis and rice, while it is poorly understood for the identity and regulatory mechanism of VPTs in Brassica napus ( B. napus). Here, we identified two vacuolar Pi influx transporters BnA09PHT5;1b and BnCnPHT5;1b in B. napus and uncovered their necessity for cellular Pi homeostasis through functional analysis. BnPHT5;1bs are the homologs of Arabidopsis AtPHT5;1 with the similar sequence, structure, tonoplast localization, and VPT activity. BnPHT5;1b double mutants had smaller shoot growth and higher shoot cellular Pi than the wild-type B. napus, which are largely different from the report in At PHT5;1 mutant, suggesting PHT5;1-VPTs play a distinct mechanism of cellular Pi homeostasis in seedlings of B. napus and Arabidopsis. By contrast, disruption of BnPHT5;1b genes slowed vegetative growth accompanied by Pi toxicity in floral organs, reduced seed yield and impacted seed traits, agreeing with the role of AtPHT5;1 in floral Pi homeostasis. Taken together, our studies identified two vacuolar Pi influx transporters in B. napus and revealed the distinct and conserved regulatory mechanisms of BnPHT5;1bs in cellular Pi homeostasis in this plant species.
The isohydric-anisohydric continuum describes the relative stringency of stomatal control of leaf water potential ( ψleaf) during drought. Hydroscape area ( HA) – the water potential landscape over which stomata regulate ψleaf – has emerged as a useful metric of the iso/anisohydric continuum because it is strongly linked to several hydraulic, photosynthetic, and structural traits. Previous research on HA focused on broad ecological patterns involving several plant clades. Here we investigate relationships of HA to climatic conditions and functional traits across ecologically diverse but closely related species while accounting for phylogeny. Across a macroclimatic moisture gradient, defined by the ratio of mean annual precipitation to mean annual pan evaporation ( P/Ep), HA decreased with P/Ep for ten Eucalyptus species. Greater anisohydry reflects lower turgor loss points and greater hydraulic safety, mirroring global patterns. More isohydric species have mesophyll photosynthetic capacity that is more sensitive to ψleaf, consistent with an earlier model for optimal stomatal behavior. Hydroscapes exhibit little plasticity in response to variation in water supply, and the extent of plasticity does not vary with P/Ep of native habitats. These findings strengthen the case that HA is a useful metric for characterizing drought tolerance and water-status regulation.