Roles for Class III HD-Zip and KANADI genes in Arabidopsis root development

Meristems within the plant body differ in their structure and the patterns and identities of organs they produce. Despite these differences, it is becoming apparent that shoot and root apical and vascular meristems share significant gene expression patterns. Class III HD-Zip genes are required for the formation of a functional shoot apical meristem. In addition, Class III HD-Zip and KANADI genes function in patterning lateral organs and vascular bundles produced from the shoot apical and vascular meristems, respectively. We utilize both gain- and loss-of-function mutants and gene expression patterns to analyze the function of Class III HD-Zip and KANADI genes in Arabidopsis roots. Here we show that both Class III HD-Zip and KANADI genes play roles in the ontogeny of lateral roots and suggest that Class III HD-Zip gene activity is required for meristematic activity in the pericycle analogous to its requirement in the shoot apical meristem.     

Carbohydrate Metabolism in Leaf Meristems of Tall Fescue : II. Relationship to Leaf Elongation Rates Modified by Nitrogen Fertilization

Our objective was to examine alterations in carbohydrate status of leaf meristems that are associated with nitrogen-induced changes in leaf elongation rates of tall fescue (Festuca arundinacea Schreb.). Dark respiration rates, concentrations of nonstructural carbohydrates, and soluble proteins were measured in leaf intercalary meristems and adjacent segments of elongating leaves. The two genotypes used differed by 43% in leaf elongation rate. Application of high nitrogen (336 kilograms per hectare) resulted in 140% higher leaf elongation rate when compared to plants receiving low nitrogen (22 kilograms per hectare). Leaf meristems of plants receiving high and low nitrogen had dark respiration rates of 5.4 and 2.9 microliters O₂ consumed per milligram structural dry weight per hour, respectively. Concentrations of soluble proteins were lower while concentrations of fructan tended to be slightly higher in leaf meristems of low-nitrogen plants when compared to high-nitrogen plants. Concentrations of reducing sugars, nonreducing sugars, and takadiastase-soluble carbohydrate of leaf meristems were not affected by nitrogen treatment. Total nonstructural carbohydrates of leaf meristems averaged 44 and 39% of dry weight for low- and high-nitrogen plants, respectively. Within the leaf meristem, approximately 74 and 34% of the pool of total nonstructural carbohydrate could be consumed per day in high- and low-nitrogen plants, respectively, assuming no carbohydrate import to the meristem occurred. Plants were able to maintain high concentrations of nonstructural carbohydrates in leaf meristems despite a 3-fold range in leaf elongation rates, suggesting that carbohydrate synthesis and transport to leaf intercalary meristems may not limit leaf growth of these genotypes.     

Study of the senescence process in primary leaves of sunflower (Helianthus annuus L.) plants under two different light intensities

Different parameters that vary during leaf development may be affected by light intensity. To study the influence of different light intensities on primary leaf senescence, sunflower (Helianthus annuus L.) plants were grown for 50 days under two photon flux density (PFD) conditions, namely high irradiance (HI) at 350 μmol(photon) m^?2 s^?1 and low irradiance (LI) at 125 μmol(photon) m^?2 s^?1. Plants grown under HI exhibited greater specific leaf mass referred to dry mass, leaf area and soluble protein at the beginning of the leaf development. This might have resulted from the increased CO₂ fixation rate observed in HI plants, during early development of primary leaves. Chlorophyll a and b contents in HI plants were lower than in LI plants in young leaves. By contrast, the carotenoid content was significantly higher in HI plants. Glucose concentration increased with the leaf age in both treatments (HI and LI), while the starch content decreased sharply in HI plants, but only slightly in LI plants. Glucose contents were higher in HI plants than in LI plants; the differences were statistically significant (p<0.05) mainly at the beginning of the leaf senescence. On the other hand, starch contents were higher in HI plants than in LI plants, throughout the whole leaf development period. Nitrate reductase (NR) activity decreased with leaf ageing in both treatments. However, the NR activation state was higher during early leaf development and decreased more markedly in senescent leaves in plants grown under HI. GS activity also decreased during sunflower leaf ageing under both PFD conditions, but HI plants showed higher GS activities than LI plants. Aminating and deaminating activities of glutamate dehydrogenase (GDH) peaked at 50 days (senescent leaves). GDH deaminating activity increased 5-fold during the leaf development in HI plants, but only 2-fold in LI plants. The plants grown under HI exhibited considerable oxidative stress in vivo during the leaf senescence, as revealed by the substantial H₂O₂ accumulation and the sharply decrease in the antioxidant enzymes, catalase and ascorbate peroxidase, in comparison with LI plants. Probably, systemic signals triggered by a high PFD caused early senescence and diminished oxidative protection in primary leaves of sunflower plants as a result.     

A Role for Glutamine Synthetase in the Remobilization of Leaf Nitrogen during Natural Senescence in Rice Leaves

Changes in the levels of cytosolic glutamine synthetase (GS1) and chloroplastic glutamine synthetase (GS2) polypeptides and of corresponding mRNAs were determined in leaves of hydroponically grown rice (Oryza sativa) plants during natural senescence. The plants were grown in the greenhouse for 105 days at which time the thirteenth leaf was fully expanded. This was counted as zero time for senescence of the twelfth leaf. The twelfth leaf blade on the main stem was analyzed over a time period of −7 days (98 days after germination) to +42 days (147 days after germination). Total GS activity declined to less than a quarter of its initial level during the senescence for 35 days and this decline was mainly caused by a decrease in the amount of GS2 polypeptide. Immunoblotting analyses showed that contents of other chloroplastic enzymes, such as ribulose-1,5-bisphosphate carboxylase/oxygenase and Fd-glutamate synthase, declined in parallel with GS2. In contrast, the GS1 polypeptide remained constant throughout the senescence period. Translatable mRNA for GS1 increased about fourfold during the senescence for 35 days. During senescence, there was a marked decrease in content of glutamate (to about one-sixth of the zero time value); glutamate is the major form of free amino acid in rice leaves. Glutamine, the major transported amino acid, increased about threefold compared to the early phase of the harvest in the senescing rice leaf blades. These observations suggest that GS1 in senescing leaf blades is responsible for the synthesis of glutamine, which is then transferred to the growing tissues in rice plants.     

HD-Zip III转录因子家族与植物细胞分化

细胞分化是生物生长发育的重要过程, 受到一系列信号的精确调控。植物特有的转录因子HD-Zip III在细胞分化中发挥了重要作用。该文对HD-Zip III基因类型和结构特点进行了简要介绍, 重点论述了HD-Zip III在胚胎形态发生、顶端分生组织形成、叶极性建立和维管组织分化等发育过程中的作用, 系统总结了HD-Zip III基因在不同层次受到的调控, 探讨了该家族基因与陆生维管植物进化的关系。     

Determination for inflorescence development is a stable state,separable from determination for flower development in Pisum sativum L. buds

Terminal meristems of Pisum sativum (garden pea) transit from vegetative to inflorescence development, and begin producing floral axillary meristems. Determination for inflorescence development was assessed by culturing excised buds and meristems. The first node of floral initiation (NFI) for bud expiants developing in culture and for adventitious shoots forming on cultured meristems was compared with the NFI of intact control buds. When terminal buds having eight leaf primordia were excised from plants of different ages (i.e., number of unfolded leaves) and cultured on 6-benzylaminopurine and kinetin-supplemented medium, the NFI was a function of the age of the source plant. By age 3, all terminal buds were determined for inflorescence development. Determination occurred at least eight nodes before the first axillary flower was initiated. Thus, the axillary meristems contributing to the inflorescence had not formed at the time the bud was explanted. Similar results were obtained for cultured axillary buds. In addition, meristems excised without leaf primordia from axillary buds three nodes above the cotyledons of age-3 plants gave rise to adventitious buds with an NFI of 8.3 ±0.3 nodes. In contrast seed-derived plants had an NFI of 16.5 ±0.2. Thus cells within the meristem were determined for inflorescence development. These findings indicate that determination for inflorescence development in P. sativum is a stable developmental state, separable from determination for flower development, and occurring prior to initiation of the inflorescence at the level of meristems.     

The Stem Cell Niche in Leaf Axils Is Established by Auxin and Cytokinin in Arabidopsis

Plants differ from most animals in their ability to initiate new cycles of growth and development, which relies on the establishment and activity of branch meristems harboring new stem cell niches. In seed plants, this is achieved by axillary meristems, which are established in the axil of each leaf base and develop into lateral branches. Here, we describe the initial processes of Arabidopsis thaliana axillary meristem initiation. Using reporter gene expression analysis, we find that axillary meristems initiate from leaf axil cells with low auxin through stereotypical stages. Consistent with this, ectopic overproduction of auxin in the leaf axil efficiently inhibits axillary meristem initiation. Furthermore, our results demonstrate that auxin efflux is required for the leaf axil auxin minimum and axillary meristem initiation. After lowering of auxin levels, a subsequent cytokinin signaling pulse is observed prior to axillary meristem initiation. Genetic analysis suggests that cytokinin perception and signaling are both required for axillary meristem initiation. Finally, we show that cytokinin overproduction in the leaf axil partially rescue axillary meristem initiation-deficient mutants. These results define a mechanistic framework for understanding axillary meristem initiation.     

GA2 and GA20-oxidase expressions are associated with the meristem position in Streptocarpus rexii (Gesneriaceae)

We examined genes involved in the regulatory pathway of gibberellin (GA) in meristems of Streptocarpus rexii. The plants do not possess a typical shoot apical meristem (SAM) and form unique meristems: the basal meristem extends the lamina area of one cotyledon to produce anisocotylous seedlings; the groove meristem forms new leaves at the base of the macrocotyledon. Exogenous application of GA significantly suppresses the basal meristem activity in developing cotyledons and the seedlings remain isocotyl. To examine the role of endogenous GA on these meristems in vivo, we isolated homologs of GA2-oxidase responsible for degrading active GAs (SrGA2ox), and GA20-oxidase regulating the rate limiting step of active GA synthesis (SrGA20ox). During embryogenesis, while first partly overlapping, the expression of SrGA2ox and SrGA20ox became more differentiated and mutually exclusive, ending with SrGA2ox being expressed solely in the adaxial–proximal domain of the embryo in regions with meristem activity, whereas SrGA20ox was restricted to the fork between the two cotyledons. The latter may be responsible for suppressing the formation of an embryonic SAM in S. rexii. In developing seedlings, SrGA2ox expression also followed the centers of meristem activity, where SrGA20ox expression was excluded. Our results suggest that low levels of GA are required in S. rexii meristems for their establishment and maintenance. Thus, the meristems in S. rexii share similar regulatory pathways suggested for the SAM in model plants, but that in S. rexii evolutionary modifications involving a lateral transfer of function, from shoot to leaves, is implicated in attaining the unusual morphology of the plants.     

Fusaric acid accelerates the senescence of leaf in banana when infected by Fusarium

Fusarium oxysporum f.sp. cubense (FOC) is a causal agent of vascular wilt and leaf chlorosis of banana plants. Chloroses resulting from FOC occur first in the lowest leaves of banana seedlings and gradually progress upward. To investigate the responses of different leaf positions to FOC infection, hydroponic experiments with FOC inoculation were conducted in a greenhouse. Fusarium-infected seedlings exhibited a decrease in net photosynthesis rate, stomatal conductance, and transpiration rate of all leaves. The wilting process in Fusarium-infected seedlings varied with leaf position. Measurements of the maximum photochemical efficiency of photosystem II (F _V/F _max) and visualization with transmission electron microscopy showed a positive correlation between chloroplast impairment and severity of disease symptoms. Furthermore, results of malondialdehyde content and relative membrane conductivity measurements demonstrated that the membrane system was damaged in infected leaves. Additionally, the activities of phenylalanine ammonia-lyase, peroxidase and polyphenol oxidase were increased and total soluble phenolic compounds were significantly accumulated in the leaves of infected plants. The structural and biochemical changes of infected plants was consistent with plant senescence. As the FOC was not detected in infected leaves, we proposed that the chloroplast and membrane could be damaged by fusaric acid produced by Fusarium. During the infection, fusaric acid was first accumulated in the lower leaves and water-soluble substances in the lower leaves could dramatically enhance fusaric acid production. Taken together, the senescence of infected banana plants was induced by Fusarium infection with fusaric acid production and the composition of different leaf positions largely contribute to the particular senescence process.     

MORPHOLOGY OF MARSILEA VESTITA. I. ONTOGENY AND MORPHOLOGY OF THE SUBMERGED AND LAND FORMS OF THE JUVENILE LEAVES

During their ontogeny, the primordia of the juvenile leaves of Marsilea plants in sterile culture develop 1, 2 or 4 marginal meristems, and these, in turn, contribute cells to the young leaf by anti- and periclinal cell divisions. The final leaves are unifid, bifid, or quadrifid, depending on how many marginal meristems develop, and this is determined early in the ontogeny of the leaf. The mechanism which determines whether or not a marginal meristem develops may fluctuate, as shown by the existence of trifid leaves. Two forms of juvenile leaves are produced, those in a liquid medium, which in many respects resemble the adult quadrifid submerged leaves, and those on a solid medium, which in many respects resemble the adult land leaves.     

Auxin Depletion from the Leaf Axil Conditions Competence for Axillary Meristem Formation in Arabidopsis and Tomato

The enormous variation in architecture of flowering plants is based to a large extent on their ability to form new axes of growth throughout their life span. Secondary growth is initiated from groups of pluripotent cells, called meristems, which are established in the axils of leaves. Such meristems form lateral organs and develop into a side shoot or a flower, depending on the developmental status of the plant and environmental conditions. The phytohormone auxin is well known to play an important role in inhibiting the outgrowth of axillary buds, a phenomenon known as apical dominance. However, the role of auxin in the process of axillary meristem formation is largely unknown. In this study, we show in the model species Arabidopsis thaliana and tomato (Solanum lycopersicum) that auxin is depleted from leaf axils during vegetative development. Disruption of polar auxin transport compromises auxin depletion from the leaf axil and axillary meristem initiation. Ectopic auxin biosynthesis in leaf axils interferes with axillary meristem formation, whereas repression of auxin signaling in polar auxin transport mutants can largely rescue their branching defects. These results strongly suggest that depletion of auxin from leaf axils is a prerequisite for axillary meristem formation during vegetative development.     

Independence of Organogenesis and Cell Pattern in Developing Angle Shoots of Selaginella martensii

Angle meristems are mounds of meristematic tissue located atdorsal and/or ventral branch points of the dichotomising stemaxes of many species of Selaginella (Lycophyta). The presentstudy examined the development of ventral angle shoots of S.martensii in response to removal of distal shoot apices (decapitation).Scanning electron microscopy of sequential replicas of developingangle meristems and angle shoots revealed that for the firsttwo pseudowhorls of leaf primordia, particular leaves are notattributable to particular merophytes of the angle meristemapical cell. Individual leaf primordia of the first (outer)pseudowhorl often form from more than one merophyte. Neitherthe shape of the angle meristem apical cell nor the directionof segmentation has any effect on the development of the angleshoot. Additionally, the apical cell of the angle meristem doesnot necessarily contribute directly to either of the new shootapices of the developing angle shoot. The first bifurcationof the angle shoot shows a remarkably consistent relationshipto the branching pattern of the parent shoot. The strong branchof the first angle shoot bifurcation typically occurs towardthe weak side branch of the parent shoot. Anatomical studiesshowed that bifurcation of the young angle shoot involved theformation of two new growth centres some distance away fromthe original angle meristem apical cell; new apical cells subsequentlyformed within these. These results provide additional supportfor the view that cell lineage has little or no effect on finalform or structure in plants.Copyright 1994, 1999 Academic Press Selaginella martensii Spring, Lycophyta, angle meristem, apical cell, shoot apical meristem, leaf primordium, branching, dichotomy, morphogenesis, determination, competence, development, mould and cast technique, replica technique, scanning electron microscopy     

Localization of axillary meristems during different stages of radish ontogenesis

An analysis of axillary meristem (axillary bud) localization of radish (Raphanus sativus L. cv. Tetra-I?ówiecka) was undertaken on vernalized (flowering) and unvernalized (vegetative) plants. It has been shown that the localization of these meristems can be different on successive nodes of the same plant and is connected with the development stages of the plants. The axillary meristems can arise on the stem as well as in the leaf axil or on the base of the subtending leaf. The localization of axillary meristems has been discussed in relation to growth directions and growth correlations inside the meristematic region of the shoot apex.     

Identification and functional characterization of a leucine-rich repeat receptor-like kinase gene that is involved in regulation of soybean leaf senescence

We report here the cloning and characterization of a soybean receptor-like kinase (RLK) gene, designated GmSARK (Glycine max senescence-associated receptor-like kinase), which is involved in regulating leaf senescence. The conceptual protein product of GmSARK contains typical domains of LRR receptor-like kinases: a cytoplasmic domain with all the 11 kinase subdomains, a transmembrane domain and an extracelullar domain containing 9 Leucine-Rich Repeat (LRR) units that may act as a receptor. The expression of GmSARK in soybean leaves was up-regulated in all the three tested senescence systems: senescing cotyledons, dark-induced primary leaf senescence and the natural leaf senescence process after florescence. Furthermore, the RNA interference (RNAi)-mediated knocking-down of GmSARK dramatically retarded soybean leaf senescence. A more complex thylakoid membrane system, higher foliar level of chlorophyll content and a very remarkable delay of senescence-induced disintegration of chloroplast structure were observed in GmSARK-RNAi transgenic leaves. A homolog of maize lethal leaf-spot 1 gene, which has been suggested to encode a key enzyme catalyzing chlorophyll breakdown, was isolated and nominated Gmlls1. The expression level of Gmgtr1 gene, which encodes a key enzyme of chlorophyll synthesis, was also analyzed. It was found that Gmlls1 was up-regulated and Gmgtr1 was down-regulated during senescence in wild-type soybean leaves. However, both of the up-regulation of Gmlls1 and down-regulation of Gmgtr1 were retarded during senescence of GmSARK-RNAi transgenic leaves. In addition, over-expression of the GmSARK gene greatly accelerated the senescence progression of CaMV 35S:GmSARK transgenic plants. Taken together, these results strongly suggested the involvement of this LRR-RLK in regulation of soybean leaf senescence, maybe via regulating chloroplast development and chlorophyll accumulation. Multiple functions of GmSARK besides its regulation of leaf senescence were also discussed. Electronic Supplementary Material Supplementary material is available for this article at Rui Gan, Peng-Li Li and Yuan-Yuan Ma contributed equally to this work.     

Leaf ontogeny of <Emphasis Type="Italic">Schinus molle</Emphasis> L. plants under cadmium contamination: the meristematic origin of leaf structural changes

Previous works show the development of thicker leaves on tolerant plants growing under cadmium (Cd²⁺) contamination. The aim of this study was to evaluate the Cd²⁺ effects on the leaf meristems of the tolerant species Schinus molle. Plants were grown in nutrient solution containing 0, 10, and 50 μM of Cd²⁺. Anatomical analysis was performed on leaf primordia sampled at regular time intervals. Under the lowest Cd²⁺ level (10 μM), increased ground meristem thickness, diameter of the cells, cell elongation rate, and leaf dry mass were found. However, 50 μM of Cd²⁺ reduced all these variables. In addition, the ground meristem cells became larger when exposed to any Cd²⁺ level. The epidermis, palisade parenchyma, and vascular tissues developed earlier in Cd²⁺-exposed leaves. The modifications found on the ground meristem may be related to the development of thicker leaves on S. molle plants exposed to low Cd²⁺ levels. Furthermore, older leaves showed higher Cd²⁺ content when compared to the younger ones, preventing the Cd²⁺ toxicity to these leaves. Thus, low Cd²⁺ concentrations change the ground meristem structure and function reflecting on the development of thicker and enhanced leaves.