The emergence of the tracheophyte-based vascular system of land plants had major impacts on the evolution of terrestrial biology, in general, through its role in facilitating the development of plants with increased stature, photosynthetic output, and ability to colonize a greatly expanded range of environmental habitats. Recently, considerable progress has been made in terms of our understanding of the developmental and physiological programs involved in the formation and function of the plant vascular system. In this review, we first examine the evolutionary events that gave rise to the tracheophytes, followed by analysis of the genetic and hormonal networks that cooperate to orchestrate vascular development in the gymnosperms and angiosperms. The two essentialfunctions performed by the vascular system, namely the delivery of resources （water, essential mineral nutrients, sugars and amino acids） to the various plant organs and provision of mechanical support are next discussed. Here, we focus on critical questions relating to structural and physiological properties controlling the delivery of material through the xylem and phloem. Recent discoveries into the role of the vascular system as an effective long-distance communication system are next assessed in terms of the coordination of developmental, physiological and defense-related processes, at the whole-plant level. A concerted effort has been made to integrate all these new findings into a comprehensive picture of the state-of-the-art in the area of plant vascular biology. Finally, areas important for future research are highlighted in terms of their likely contribution both to basic knowledge and applications to primary industry.
CRISPR/Cas9 uses a guide RNA （gRNA） molecule to execute sequence-specific DNA cleavage and it has been widely used for genome editing in many organisms. Modifications at either end of the gRNAs often render Cas9/gRNA inactive. So far, production of gRNA in vivo has only been achieved by using the U6 and U3 snRNA promoters. However, the U6 and U3 promoters have major limitations such as a lack of cell specificity and unsuitability for in vitro transcription. Here, we present a versatile method for efficiently producing gRNAs both in vitro and in vivo. We design an artificial gene named RGR that, once transcribed, generates an RNA molecule with ribozyme sequences at both ends of the designed gRNA. We show that the primary transcripts of RGR undergo self-catalyzed cleavage to generate the desired gRNA, which can efficiently guide sequence-specific cleavage of DNA targets both in vitro and in yeast. RGR can be transcribed from any promoters and thus allows for cell- and tissue-specific genome editing if appropriate promoters are chosen. Detecting mutations generated by CRISPR is often achieved by enzyme digestions, which are not very compatible with high-throughput analysis. Our system allows for the use of universal primers to produce any gRNAs in vitro, which can then be used with Cas9 protein to detect mutations caused by the gRNAs/CRISPR. In conclusion, we provide a versatile method for generating targeted mutations in specific cells and tissues, and for efficiently detecting the mutations generated.
Lignin is a polymer of phenylpropanoid compounds formed through a complex biosynthesis route, represented by a metabolic grid for which most of the genes involved have been sequenced in several plants, mainly in the model-plants Arabidopsis thaliana and Populus. Plants are exposed to different stresses, which may change lignin content and composition. In many cases, particularly for plant-microbe interactions, this has been suggested as defence responses of plants to the stress. Thus, understanding how a stressor modulates expression of the genes related with lignin biosynthesis may allow us to develop study-models to increase our knowledge on the metabolic control of lignin deposition in the cell wall. This review focuses on recent literature reporting on the main types of abiotic and biotic stresses that alter the biosynthesis of lignin in plants.
As an essential plant macronutrient, the low availability of phosphorus （P） in most soils imposes serious limitation on crop production. Plants have evolved complex responsive and adaptive mechanisms for acquisition, remobilization and recycling of phosphate （Pi） to maintain P homeostasis. Spatio-temporal molecular, physiological, and biochemical Pi deficiency responses developed by plants are the consequence of local and systemic sensing and signaling pathways. Pi deficiency is sensed locally by the root system where hormones serve as important signaling components in terms of developmental reprogramming, leading to changes in root system architecture. Root-to-shoot and shoot-to-root signals, delivered through the xylem and phloem, respectively, involving Pi itself, hormones, miRNAs, mRNAs, and sucrose, serve to coordinate Pi deficiency responses at the whole-plant level. A combination of chromatin remodeling, transcriptional and posttranslational events contribute to globally regulating a wide range of Pi deficiency responses. In this review, recent advances are evaluated in terms of progress toward developing a comprehen- sive understanding of the molecular events underlying control over P homeostasis. Application of this knowledge, in terms of developing crop plants having enhanced attributes for P use efficiency, is discussed from the perspective of agricultural sustainability in the face of diminishing global P supplies.
Flavonoids are ubiquitous in the plant kingdom and have many diverse functions including defense, UV protection, auxin transport inhibition, allelopathy, and flower coloring. Interestingly, these compounds also have considerable biological activity in plant, animal and bacterial systems - such broad activity is accomplished by few compounds. Yet, for all the research over the last three decades, many of the cellular targets of these secondary metabolites are unknown. The many mutants available in model plant species such as Arabidopsis thaliana and Medicago truncatula are enabling the intricacies of the physiology of these compounds to be deduced. In the present review, we cover recent advances in flavonoid research, discuss deficiencies in our understanding of the physiological processes, and suggest approaches to identify the cellular targets of flavonoids.
Legumes are highly important food, feed and biofuel crops. With few exceptions, they can enter into an intricate symbiotic relationship with specific soil bacteria called rhizobia. This interaction results in the formation of a new root organ called the nodule in which the rhizobia convert atmospheric nitrogen gas into forms of nitrogen that are useable by the plant. The plant tightly controls the number of nodules it forms, via a complex root-to-shoot-to-root signaling loop called autoregulation of nodulation （AON）. This regulatory process involves peptide hormones, receptor kinases and small metabolites. Using modern genetic and genomic techniques, many of the components required for nodule formation and AON have now been isolated. This review addresses these recent findings, presents detailed models of the nodulation and AON processes, and identifies gaps in our understanding of these process that have yet to be fully explained.
The small phenolic compound salicylic acid （SA） plays an important regulatory role in multiple physiological processes including plant im- mune response. Significant progress has been made during the past two decades in understanding the SA-mediated defense signaling network. Characterization of a number of genes functioning in SA biosynthesis, conjugation, accumulation, signaling, and crosstalk with other hormones such as jasmonic acid, ethylene, abscisic acid, auxin, gibberellic acid, cytokinin, brassinosteroid, and peptide hormones has sketched the finely tuned immune response network. Full understanding of the mech- anism of plant immunity will need to take advantage of fast developing genomics tools and bioinformatics techniques. However, elucidating genetic components involved in these pathways by conventional ge- netics, biochemistry, and molecular biology approaches will continue to be a major task of the community. High-throughput method for SA quantification holds the potential for isolating additional mutants related to SA-mediated defense signaling.
You‐Zhi Ma (Corresponding author) Plants have acquired sophisticated stress response systems to adapt to changing environments. It is important to understand plants’ stress response mechanisms in the effort to improve crop productivity under stressful conditions. The AP2/ERF transcription factors are known to regulate diverse processes of plant development and stress responses. In this study, the molecular characteristics and biological functions of AP2/ERFs in a variety of plant species were analyzed. AP2/ERFs, especially those in DREB and ERF subfamilies, are ideal candidates for crop improvement because their overexpression enhances tolerances to drought, salt, freezing, as well as resistances to multiple diseases in the transgenic plants. The comprehensive analysis of physiological functions is useful in elucidating the biological roles of AP2/ERF family genes in gene interaction, pathway regulation, and defense response under stress environments, which should provide new opportunities for the crop tolerance engineering.
The WRKY gene family is among the largest families of transcription factors （TFs） in higher plants. By regulating the plant hormone signal transduction pathway, these TFs play critical roles in some plant processes in response to biotic and abiotic stress, Various bodies of research have demonstrated the important biological functions of WRKY TFs in plant response to different kinds of biotic and abiotic stresses and working mecha- nisms. However, very li2ttle summarization has been done to review their research progress. Not iust important TFs function in plant response to biotic and abiotic stresses, WRKY also participates in carbohydrate synthesis, senes- cence, development, and secondary metabolites synthesis. WRKY proteins can bind to W-box （TGACC （A/T）） in the promoter of its target genes and activate or repress the expression of downstream genes to regulate their stress response. Moreover, WRKY proteins can interact with other TFs to regulate plant defensive responses. In the present review, we focus on the structural characteristics of WRKY TFs and the research progress on their functions in plant responses to a variety of stresses.
Genomic selection （GS） and high-throughput phenotyping have recently been captivating the interest of the crop breeding com- munity from both the public and private sectors world-wide. Both approaches promise to revolutionize the prediction of complex traits, including growth, yield and adaptation to stress. Whereas high-throughput phenotyping may help to improve understanding of crop physiology, most powerful techniques for high-throughput field phenotyping are empirical rather than analytical and compa- rable to genomic selection. Despite the fact that the two method- ological approaches represent the extremes of what is understood as the breeding process （phenotype versus genome）, they both consider the targeted traits （e.g. grain yield, growth, phenology, plant adaptation to stress） as a black box instead of dissectingthem as a set of secondary traits （i.e. physiological） putatively related to the target trait. Both GS and high-throughput phenotyping have in common their empirical approach enabling breeders to use genome profile or phenotype without understanding the underlying biology. This short review discusses the main aspects of both approaches and focuses on the case of genomic selection of maize flowering traits and near-infrared spectroscopy （NIRS） and plant spectral reflectance as high-throughput field phenotyping methods for complex traits such as crop growth and yield.
Pollen development is a critical step in plant development that is needed for successful breeding and seed formation. Manipulation of male fertility has proved a useful trait for hybrid breeding and increased crop yield. However, although there is a good understanding developing of the molecular mechanisms of anther and pollen anther development in model species, such as Arabidopsis and rice, little is known about the equivalent processes in important crops. Nevertheless the onset of increased genomic information and genetic tools is facilitating translation of information from the models to crops, such as barley and wheat; this will enable increased understanding and manipulation of these pathways for agricultural improvement.
Potassium （K＋） is an essential macronutrient in plants and a lack of K＋ significantly reduces the potential for plant growth and development. By contrast, sodium （Na＋）, while beneficial to some extent, at high concentrations it disturbs and inhibits various physiological processes and plant growth. Due to their chemical similarities, some functions of K＋ can be undertaken by Na＋ but K＋ homeostasis is severely affected by salt stress, on the other hand. Recent advances have highlighted the fascinating regulatory mechanisms of K＋ and Na＋ transport and signaling in plants. This review summarizes three major topics： （i） the transport mechanisms of K＋ and Na＋ from the soil to the shoot and to the cellular - compartments; （ii） the mechanisms through which plants sense and respond to K＋ and Na＋ availability; and （iii） the components involved in maintenance of K＋/Na＋ homeostasis in plants under salt stress.
Plant secondary metabolites play critical roles in plant-environment interactions. They are synthesized in different organs or tissues at particular developmental stages, and in response to various environmental stimuli, both biotic and abiotic. Accordingly, corresponding genes are regulated at the transcriptional level by multiple transcription factors. Several families of transcription factors have been identified to participate in controlling the biosynthesis and accumulation of secondary metabolites. These regulators integrate internal （often developmental） and external signals, bind to corresponding cis-elements which are often in the promoter regions to activate or repress the expression of enzyme-coding genes, and some of them interact with other transcription factors to form a complex. In this review, we summarize recent research in these areas, with an emphasis on newly-identified transcription factors and their functions in metabolism regulation.
Reactive Oxygen Species （ROS） are continuously produced as a result of aerobic metabolism or in response to biotic and abiotic stresses. ROS are not only toxic by-products of aerobic metabolism, but are also signalling molecules involved in several developmental processes in all organisms. Previous studies have clearly shown that an oxidative burst often takes place at the site of attempted invasion during the early stages of most plant-pathogen interac- tions. Moreover, a second ROS production can be observed during certain types of plant-pathogen interactions, which triggers hyper- sensitive cell death （HR）. This second ROS wave seems absent during symbiotic interactions. This difference between these two responses is thought to play an important signalling role leading to the establishment of plant defense. In order to cope with the deleterious effects of ROS, plants are fitted with a large panel of enzymatic and non-enzymatic antioxidant mechanisms. Thus,increasing numbers of publications report the characterisation of ROS producing and scavenging systems from plants and from microorganisms during interactions. In this review, we present the current knowledge on the ROS signals and their role during plant-microorganism interactions.
Polyamines （mainly putrescine （Put）, spermidine （Spd）, and spermine （Spin）） have been widely found in a range of physiological processes and in almost all diverse environ- mental stresses. In various plant species, abiotic stresses modulated the accumulation of polyamines and related gene expression. Studies using loss-of-function mutants and transgenic overexpression plants modulating polyamine metabolic pathways confirmed protective roles of polyamines during plant abiotic stress responses, and indicated the possibility to improve plant tolerance through genetic manipulation of the polyamine pathway. Additionally, puta- tive mechanisms of polyamines involved in plant abiotic stress tolerance were thoroughly discussed and crosstalks among polyamine, abscisic acid, and nitric oxide in plant responses to abiotic stress were emphasized. Special attention was paid to the interaction between polyamine and reactive oxygen species, ion channels, amino acid and carbon metabolism, and other adaptive responses. Further studies are needed to elucidate the polyamine signaling pathway, especially polyamine-regulated downstream tar- gets and the connections between polyamines and other stress responsive molecules.
Water is transported in xylem conduits from the soil to the leaves under negative pressures. This is a metabolically inexpensive transport system but it is subjected to the risk of cavitation failure. Herein we address methodologies for measuring xylem resistance to cavitation, current debates, and future research opportunities in plant hydraulics.
The productivity, product quality and competitive ability of important agricultural and horticultural plants in many regions of the world may be adversely affected by current and anticipated concentrations of ground-level ozone (O-3). Exposure to elevated O-3 typically results in suppressed photosynthesis, accelerated senescence, decreased growth and lower yields. Various approaches used to evaluate O-3 effects generally concur that current yield losses range from 5% to 15% among sensitive plants. There is, however, considerable genetic variability in plant responses to O-3. To illustrate this, we show that ambient O-3 concentrations in the eastern United States cause substantially different levels of damage to otherwise similar snap bean cultivars. Largely undesirable effects of O-3 can also occur in seed and fruit chemistry as well as in forage nutritive value, with consequences for animal production. Ozone may alter herbicide efficacy and foster establishment of some invasive species. We conclude that current and projected levels of O-3 in many regions worldwide are toxic to sensitive plants of agricultural and horticultural significance. Plant breeding that incorporates O-3 sensitivity into selection strategies will be increasingly necessary to achieve sustainable production with changing atmospheric composition, while reductions in O-3 precursor emissions will likely benefit world food production and reduce atmospheric concentrations of an important greenhouse gas.
Citrus huanglongbing (HLB) is the most important disease in the citrus industry. HLB development and severity are a function of multiple elements, in which plant innate immunity, pathogen population in a specific growth region, vector spreading and jumping between regions and among hosts are major contributors.
With the enhancement of copper (Cu) stress, the germination percentage of wheat seeds decreased gradually. Pretreatment with sodium hydrosulfide (NaHS), hydrogen sulfide (H2S) donor alleviated the inhibitory effect of Cu stress in a dose-dependent manner; whereas little visible symptom was observed in germinating seeds and radicle tips cultured in NaHS solutions. It was verified that H2S or HS- rather than other sulfur-containing components derived from NaHS attribute to the potential role in promoting seed germination against Cu stress. Further studies showed that NaHS could promote amylase and esterase activities, reduce Cu-induced disturbance of plasma membrane integrity in the radicle tips, and sustain lower levels of malondialdehyde and H2O2 in germinating seeds. Furthermore, NaHS pretreatment increased activities of superoxide dismutase and catalase and decreased that of lipoxygenase, but showed no significant effect on ascorbate peroxidase. Alternatively, NaHS prevented uptake of Cu and promoted the accumulation of free amino acids in seeds exposed to Cu. In addition, a rapid accumulation of endogenous H2S in seeds was observed at the early stage of germination, and higher level of H2S in NaHS-pretreated seeds. These data indicated that H2S was involved in the mechanism of germinating seeds' responses to Cu stress.
As researchers have gained a better understanding in recent years into the physiological, molecular, and genetic basis of how plants deal with aluminum （AI） toxicity in acid soils prevalent in the tropics and sub-tropics, it has become clear that an important component of these responses is the triggering and regulation of cellular pathways and processes by AI. In this review of plant AI signaling, we begin by summarizing the understanding of physiological mechanisms of AI resistance, which first led researchers to realize that AI stress induces gene expression and modifies protein function during the activation of AI resistance responses. Subsequently, an overview of AI resistance genes and their function provides verification that AI induction of gene expression plays a major role in AI resistance in many plant species. More recent research into the mechanistic basis for Al-induced transcrip- tional activation of resistance genes has led to the identifica- tion of several transcription factors as well as cis-elements in the promoters of AI resistance genes that play a role in greater Al-induced gene expression as well as higher constitutive expression of resistance genes in some plant species. Finally, the post-transcriptional and translational regulation of AI resistance proteins is addressed, where recent research has shown that AI can both directly bind to and alter activity of certain organic acid transporters, and also influence AI resistance proteins indirectly, via protein phosphorylation.