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PubMed1996

The WUSCHEL gene is required for shoot and floral meristem integrity in Arabidopsis.

Self perpetuation of the shoot meristem is essential for the repetitive initiation of shoot structures during plant development. In Arabidopsis shoot meristem maintenance is disrupted by recessive mutations in the WUSCHEL (WUS) gene. The defect is evident at all developmental stages and is restricted to shoot and floral meristems, whereas the root meristem is not affected. wus mutants fail to properly organize a shoot meristem in the embryo. Postembryonically, defective shoot meristems are initiated repetitively but terminate prematurely in aberrant flat structures. In contrast to wild-type shoot meristems, primordia initiation occurs ectopically across mutant apices, including the center, and often new shoot meristems instead of organs are initiated. The cells of wus shoot apices are larger and more vacuolated than wild-type shoot meristem cells. wus floral meristems terminate prematurely in a central stamen. Double mutant studies indicate that the number of organ primordia in the center of wus flowers is limited, irrespective of organ identity and we propose that meristem cells are allocated into floral whorl domains in a sequential manner. WUS activity also appears to be required for the formation of supernumerary organs in the center of agamous, superman or clavata1 flowers, suggesting that the WUS gene acts upstream of the corresponding genes. Our results suggest that the WUS gene is specifically required for central meristem identity of shoot and floral meristems to maintain their structural and functional integrity.

Arabidopsis↗

PubMed1998

The control of maize spikelet meristem fate by the APETALA2-like gene indeterminate spikelet1.

The orderly production of meristems with specific fates is crucial for the proper elaboration of plant architecture. The maize inflorescence meristem branches several times to produce lateral meristems with determinate fates. The first meristem formed, the spikelet pair meristem, produces two spikelet meristems, each of which produces two floral meristems. We have identified a gene called indeterminate spikelet1 (ids1) that specifies a determinate spikelet meristem fate and thereby limits the number of floral meristems produced. In the absence of ids1 gene function, the spikelet meristem becomes indeterminate and produces additional florets. Members of the grass family vary in the number of florets within their spikelets, suggesting that ids1 may play a role in inflorescence architecture in other grass species. ids1 is a member of the APETALA2 (AP2) gene family of transcription factors that has been implicated in a wide range of plant development roles. Expression of ids1 was detected in many types of lateral organ primordia as well as spikelet meristems. Our analysis of the ids1 mutant phenotype and expression pattern indicates that ids1 specifies determinate fates by suppressing indeterminate growth within the spikelet meristem.

Amino Acid Sequence↗

PubMed2003

The DORNROSCHEN/ENHANCER OF SHOOT REGENERATION1 gene of Arabidopsis acts in the control of meristem ccll fate and lateral organ development.

The two main tasks of a meristem, self-perpetuation and organ initiation, are separated spatially. Slowly dividing cells in the meristem center act as pluripotent stem cells, and only their derivatives in the meristem periphery specify new organs. Meristem integrity and cellular proliferation are controlled in part by regulatory interactions between genes that are expressed in specific subdomains of the meristem. Using transposon-mediated activation tagging, we have identified Dornröschen (drn-D) mutants of Arabidopsis that prematurely arrest shoot meristem activity with the formation of radialized lateral organs. The mutated gene (DRN/ESR1), which encodes an AP2/ERF protein, is expressed in a subdomain of meristem stem cells, in lateral organ anlagen, and transiently in the distal domain of organ primordia. During the development of drn-D mutants, expression of the homeobox gene SHOOTMERISTEMLESS is downregulated and later reactivated in an altered domain. In addition, we found increased expression of CLAVATA3 and WUSCHEL, two genes that antagonistically regulate stem cell fate in meristems. These findings suggest that the DRN/ESR1 gene product is involved in the regulation of gene expression patterns in meristems. Furthermore, specific misexpression of DRN in meristem stem cells affects organ polarity and outgrowth in the meristem periphery, indicating that DRN/ESR1 itself, or a process regulated by DRN/ESR1, can act non-cell-autonomously. We elaborate on the role of DRN/ESR1 in meristem and organ development and discuss its possible role in the process of shoot regeneration.

Alleles↗

PubMed1998

Cellular parameters of the shoot apical meristem in Arabidopsis.

The shoot apical meristem (SAM) is a small group of dividing cells that generate all of the aerial parts of the plant. With the goal of providing a framework for the analysis of Arabidopsis meristems at the cellular level, we performed a detailed morphometric study of actively growing inflorescence apices of the Landsberg erecta and Wassilewskija ecotypes. For this purpose, cell size, spatial distribution of mitotic cells, and the mitotic index were determined in a series of optical sections made with a confocal laser scanning microscope. The results allowed us to identify zones within the inflorescence SAM with different cell proliferation rates. In particular, we were able to define a central area that was four to six cells wide and had a low mitotic index. We used this technique to compare the meristem of the wild type with the enlarged meristems of two mutants, clavata3-1 (clv3-1) and mgoun2 (mgo2). One of the proposed functions of the CLV genes is to limit cell division rates in the center of the meristem. Our data allowed us to reject this hypothesis, because the mitotic index was reduced in the inflorescence meristem of the clv3-1 mutant. We also observed a large zone of slowly dividing cells in meristems of clv3-1 seedlings. This zone was not detectable in the wild type. These results suggest that the central area is increased in size in the mutant meristem, which is in line with the hypothesis that the CLV3 gene is necessary for the transition of cells from the central to the peripheral zone. Genetic and microscopic analyses suggest that mgo2 is impaired in the production of primordia, and we previously proposed that the increased size of the mgo2 meristem could be due to an accumulation of cells at the periphery. Our morphometric analysis showed that mgo2 meristems, in contrast to those of clv3-1, have an enlarged periphery with high cell proliferation rates. This confirms that clv3-1 and mgo2 lead to meristem overgrowth by affecting different aspects of meristem function.

Arabidopsis↗

PubMed1994

Cell-layer interactions influence the number and position of lateral shoot meristems in Nicotiana.

In higher plants the formation of lateral shoot meristems (i.e., axillary meristems) in the axils of leaves establishes potential growth centers along the principal axis of the stem. The position and number of lateral buds in Nicotiana glauca, two genotypes of Nicotiana tabacum, and a series of interspecific periclinal chimeras composed of these species was studied to establish the role of position (location of the node along the main axis), flowering, and cell-layer interactions on the pattern of lateral meristem initiation. In N. glauca, both the number of nodes generated and the transition to flowering influenced the number and position of lateral meristems. A short-day mutant of N. tabacum grown under long days remained vegetative and never produced multiple lateral buds per node, indicating that attaining a certain node number was not sufficient to cause the formation of multiple buds. Yet, flowering plants of both short-day and day-neutral N. tabacum possessed multiple buds in their upper nodes. An analysis of the number of buds per node and bud position along the main axis in periclinal chimeras indicated that the genotype of the third apical layer (LIII) of the meristem had the greatest influence on the pattern of lateral shoot meristems in both vegetative and flowering plants. The lineage of the three apical layers (LI, LII, and LIII) of the terminal shoot meristem was preserved in primary (1 degree) lateral meristems but minor deviations in lineage were observed in secondary (2 degrees) buds (i.e., those formed later but in the same node as the 1 degree bud). An analysis of the phenotype of 2 degrees shoots that displayed deviations from expected lineage indicated that in most cases the periclinal cell divisions that destabilized the lineage occurred at the flanks of the meristem and began before the most basal node, indicating that periclinal cell divisions most likely occurred prior to the inception of the 2 degrees lateral meristem. Based on our studies, we conclude that both 1 degree and 2 degrees lateral meristems in Nicotiana ultimately descend from derivatives of all three apical layers of the terminal shoot meristem.

Plant Cells↗

PubMed2003

ZWILLE buffers meristem stability in Arabidopsis thaliana.

The shoot apical meristem of higher plants consists of a population of stem cells at the tip of the plant body that continuously gives rise to organs such as leaves and flowers. Cells that leave the meristem differentiate and must be replaced to maintain the integrity of the meristem. The balance between differentiation and maintenance is governed both by the environment and the developmental status of the plant. In order to respond to these different stimuli, the meristem has to be plastic thus ensuring the stereotypic shape of the plant body. Meristem plasticity requires the ZWILLE (ZLL) gene. In zll mutant embryos, the apical cells are misspecified causing a variability of the meristem's size and function. Using specific antibodies against ZLL, we show that the zll phenotype is due to the complete absence of the ZLL protein. In immunohistochemical experiments we confirm the observation that ZLL is solely localized in vascular tissue. For a better understanding of the role of ZLL in meristem stability, we analysed the genetic interactions of ZLL with WUSCHEL (WUS) and the CLAVATA1, 2 and 3 (CLV) genes that are involved in size regulation of the meristem. In a zll loss-of-function background wus has a negative effect whereas clv mutations have a positive effect on meristem size. We propose that ZLL buffers meristem stability non-cell-autonomously by ensuring the critical number of apical cells required for proper meristem function.

Alleles↗

PubMed2006

Nutrient sensing in plant meristems.

Plants need nutrient to grow and plant cells need nutrient to divide. The meristems are the factories and cells that are left behind will expand and differentiate. However, meristems are not simple homogenous entities; cells in different parts of the meristem do different things. Positional cues operate that can fate cells into different tissue domains. However, founder/stem cells persist in specific locations within the meristem e.g. the quiescent centre of root apical meristem (RAM) and the lower half of the central zone of the shoot apical meristem (SAM). Given the complexity of meristems, do their cells simply respond to a diffusing gradient of photosynthate? This in turn begs the question, why do stem cell populations tend to have longer cell cycles than their immediate descendants given that like all other cells they are directly in the path of diffusing nutrient? In this review, we have examined the extent to which nutrient sensing might be operating in meristems. The scene is set for sugar sensing, the plant cell cycle, SAMs and RAMs. Special emphasis is given to the metabolic regulator, SnRK1 (SNF1-related protein kinase 1), hexokinase and the trehalose pathway in relation to sugar sensing. The unique plant cell cycle gene, cyclin-dependent kinase B1;1 may have evolved to be particularly responsive to sugar signalling pathways. Also, the homeobox gene, STIMPY, emerges strongly as a link between sugar sensing, plant cell proliferation and development. Flowering can be influenced by sucrose and glucose levels and both meristem identity and organ identity genes could well be differentially sensitive to sucrose and glucose signals. We also describe how meristems deal with extra photosynthate as a result of exposure to elevated CO2. What we review are numerous instances of how developmental processes can be affected by sugars/nutrients. However, given the scarcity of knowledge we are unable to provide uncontested links between nutrient sensing and specific activities in meristems.

Carbohydrate Metabolism↗

PubMed1984

Carbohydrate metabolism in leaf meristems of tall fescue : I. Relationship to genetically altered leaf elongation rates.

The physiological bases for genetic differences in leaf growth rates were examined in two genotypes of tall fescue (Festuca arundinacea Schreb.) selected for a 50% difference in leaf elongation rate. Genotypes had similar dark respiration rates and concentrations of carbohydrate fractions in the leaf meristem and in each daily growth segment above the meristem. Dark respiration rates and concentrations of nonreducing sugars, fructans, and takadiastase-soluble carbohydrates were highest in leaf intercalary meristems and declined acropetally with tissue age. Concentrations of reducing sugars were 1.0% of dry weight in leaf meristems, 3.7% of dry weight in tissue adjacent to the meristem, then decreased progressively with distance from the meristem. Glucose, fructose, and myo-inositol comprised over 90% of the monosaccharides present in leaf meristems. Soluble protein concentration was 9.7 milligrams per gram fresh weight in leaf meristems, 5.5 milligrams per gram in tissues immediately above the meristem and, thereafter, increased linearly with distance from the meristem.Leaf meristems of the genotype exhibiting rapid leaf elongation contained 30% more soluble protein than those of the genotype selected for slow leaf elongation. The 4-fold difference in size of the leaf meristem appeared to be more important in influencing leaf elongation than were other characteristics examined.

Journal Article↗

PubMed2001

REVOLUTA regulates meristem initiation at lateral positions.

While the shoot apical meristem (SAM) is indirectly responsible for the initiation of all above-ground postembryonic organs, in most plants the vast majority of these organs are directly initiated by lateral meristems. In Arabidopsis thaliana, the lateral meristems include flower meristems (FMs), which form on the flanks of the SAM, and lateral shoot meristems (LSMs), which develop in leaf axils. While significant progress has been made on the molecular genetic basis of SAM initiation during embryo development, relatively little is known about the initiation of meristems at lateral positions. Here we have characterized the phenotypic consequences and genetic interactions of mutations in the REVOLUTA (REV) gene, with an emphasis on the role of REV in lateral meristem initiation. Our observations indicate that REV is required for initiation of both LSMs and FMs, and likely acts in the same pathway as, and upstream of, known meristem regulators. We identified the REV gene and found it encodes a predicted homeodomain/leucine zipper transcription factor that also contains a START sterol-lipid binding domain. REV is the same as the IFL gene. REV was expressed at the earliest stages of LSM and FM formation. Within the inflorescence shoot meristem, REV expression appeared to predict 3--5 incipient flower primordia on the flanks of the SAM, and REV expression at stage 1 and stage 2 matched that of WUS and STM, respectively. We propose that REV acts at lateral positions to activate the expression of known meristem regulators.

Arabidopsis↗

PubMed2004

The gene FLORAL ORGAN NUMBER1 regulates floral meristem size in rice and encodes a leucine-rich repeat receptor kinase orthologous to Arabidopsis CLAVATA1.

The regulation of floral organ number is closely associated with floral meristem size. Mutations in the gene FLORAL ORGAN NUMBER1 (FON1) cause enlargement of the floral meristem in Oryza sativa (rice), resulting in an increase in the number of all floral organs. Ectopic floral organs develop in the whorl of each organ and/or in the additional whorls that form. Inner floral organs are more severely affected than outer floral organs. Many carpel primordia develop indeterminately, and undifferentiated meristematic tissues remain in the center in almost-mature flowers. Consistent with this result, OSH1, a molecular marker of meristematic indeterminate cells in rice, continues to be expressed in this region. Although floral meristems are strongly affected by the fon1-2 mutation, vegetative and inflorescence meristems are largely normal, even in this strong allele. We isolated the FON1 gene by positional cloning and found that it encodes a leucine-rich repeat receptor-like kinase most similar to CLAVATA1 (CLV1) in Arabidopsis thaliana. This suggests that a pathway similar to the CLV signaling system that regulates meristem maintenance in Arabidopsis is conserved in the grass family. Unlike CLV1, which is predominantly expressed in the L3 layer of the shoot meristem, FON1 is expressed throughout the whole floral meristem, suggesting that small modifications to the CLV signaling pathway may be required to maintain the floral meristem in rice. In addition, FON1 transcripts are detected in all meristems responsible for development of the aerial part of rice, suggesting that genes sharing functional redundancy with FON1 act in the vegetative and inflorescence meristems to mask the effects of the fon1 mutation.

Amino Acid Sequence↗

PubMed2005

ULTRAPETALA1 encodes a SAND domain putative transcriptional regulator that controls shoot and floral meristem activity in Arabidopsis.

The higher-plant shoot apical meristem is a dynamic structure continuously producing cells that become incorporated into new leaves, stems and flowers. The maintenance of a constant flow of cells through the meristem depends on coordination of two antagonistic processes: self-renewal of the stem cell population and initiation of the lateral organs. This coordination is stringently controlled by gene networks that contain both positive and negative components. We have previously defined the ULTRAPETALA1 (ULT1) gene as a key negative regulator of cell accumulation in Arabidopsis shoot and floral meristems, because mutations in ULT1 cause the enlargement of inflorescence and floral meristems, the production of supernumerary flowers and floral organs, and a delay in floral meristem termination. Here, we show that ULT1 negatively regulates the size of the WUSCHEL (WUS)-expressing organizing center in inflorescence meristems. We have cloned the ULT1 gene and find that it encodes a small protein containing a B-box-like motif and a SAND domain, a DNA-binding motif previously reported only in animal transcription factors. ULT1 and its Arabidopsis paralog ULT2 define a novel small gene family in plants. ULT1 and ULT2 are expressed coordinately in embryonic shoot apical meristems, in inflorescence and floral meristems, and in developing stamens, carpels and ovules. Additionally, ULT1 is expressed in vegetative meristems and leaf primordia. ULT2 protein can compensate for mutant ULT1 protein when overexpressed in an ult1 background, indicating that the two genes may regulate a common set of targets during plant development. Downregulation of both ULT genes can lead to shoot apical meristem arrest shortly after germination, revealing a requirement for ULT activity in early development.

Alleles↗

PubMed2001

The ULTRAPETALA gene controls shoot and floral meristem size in Arabidopsis.

The regulation of proper shoot and floral meristem size during plant development is mediated by a complex interaction of stem cell promoting and restricting factors. The phenotypic effects of mutations in the ULTRAPETALA gene, which is required to control shoot and floral meristem cell accumulation in Arabidopsis thaliana, are described. ultrapetala flowers contain more floral organs and whorls than wild-type plants, phenotypes that correlate with an increase in floral meristem size preceding organ initiation. ultrapetala plants also produce more floral meristems than wild-type plants, correlating with an increase in inflorescence meristem size without visible fasciation. Expression analysis indicates that ULTRAPETALA controls meristem cell accumulation partly by limiting the domain of CLAVATA1 expression. Genetic studies show that ULTRAPETALA acts independently of ERA1, but has overlapping functions with PERIANTHIA and the CLAVATA signal transduction pathway in controlling shoot and floral meristem size and meristem determinacy. Thus ULTRAPETALA defines a novel locus that restricts meristem cell accumulation in Arabidopsis shoot and floral meristems.

Arabidopsis↗

PubMed1984

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 O(2) 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.

Journal Article↗

PubMed2000

Meristem growth dynamics and branching patterns in the Cladoniaceae.

Branching patterns in the lichen family Cladoniaceae are varied and taxonomically important. Branching occurs on the podetium, the erect secondary thallus that characterizes most species in the Cladoniaceae, and is influenced by growth dynamics of the fungal meristem tissue at the apex of the podetium. Branching is primarily the result of meristem divisions, and branching patterns are modified by meristem enlargement, deformation, and torsion. Branching processes are conserved, and early branch ontogeny provides information from which to determine relationships in the Cladoniaceae. Branching is characterized by two major patterns. In one pattern, branches arise from the relatively late divisions of a large meristem (≥100 μm in diameter), whose shape changes during ontogeny. In a second pattern, branches arise from small meristems (<100 μm in diameter), which split early in ontogeny but whose shape does not change. The trend toward reduced meristems that split early in ontogeny is seen as an evolutionary advance in the Cladoniaceae. Some "small meristem" species retain aspects of the "large meristem" habit in early ontogeny, and this provides a clue to their relationships. Patterns of meristem growth dynamics provide a basis for interpreting phylogeny in mycobionts of the Cladoniaceae. Meristem activities in four genera of the Cladoniaceae were compared in order to determine trends in growth dynamics within the family.

Journal Article↗

PubMed2001

Functional domains in plant shoot meristems.

The development of higher plants depends on the activity of a shoot apical meristem. Organs are formed on the flanks of the meristem, while pluripotent stem cells are found in a separate domain in the meristem centre. Further domains are distinguished by the expression patterns of genes that control the development of the shoot meristem. Although most plant cells are immobile, their relative position within a meristem, and therefore also their function, can change after cell divisions. To maintain an active shoot meristem throughout plant life, the cells in the meristem need constantly to assess their position, transmit this information to others, and readjust their gene expression profiles and their fate. Some of the genes that permit intercellular communication have been isolated. They enable the flow of information in and between meristem regions via ligands and receptor proteins to transcription factors, that ultimately control the fate of cells in the centre of the meristem.

Animals↗

PubMed2002

Axillary meristem development in the branchless Zu-0 ecotype of Arabidopsis thaliana.

Axillary meristems form in the leaf axils during post-embryonic development. In order to initiate the genetic dissection of axillary meristem development, we have characterized the late-flowering branchless ecotype of Arabidopsis thaliana (L.) Heynh., Zu-0. The first-formed rosette leaves of Zu-0 plants all initiate axillary meristems, but later-formed leaves of the rosette remain branchless. Alteration in the meristem development is axillary meristem-specific because the shoot apical and floral meristems develop normally. Scanning electron microscopy, histology and RNA in situ analysis with SHOOTMERISTEMLESS ( STM), a marker for meristematic tissues, show that a mound of cells form and STM mRNA accumulates in barren leaf axils, indicating that axillary meristems initiate but arrest in their development prior to organizing a meristem proper. Expression and retention of the STM RNA in barren leaf axils further suggests that STM expression is not sufficient for the establishment of the axillary meristem proper.

Arabidopsis↗

PubMed2003

The SHOOTLESS2 and SHOOTLESS1 genes are involved in both initiation and maintenance of the shoot apical meristem through regulating the number of indeterminate cells.

To characterize the SHL2 and SHL1 genes in detail, we analyzed three strains carrying weak alleles of SHL2, shl2-6, shl2-7, and shl2-8, and one weak allele of SHL1, shl1-3. In contrast to strong alleles, which result in lack of shoot meristem, strains bearing these weak alleles formed shoot meristem frequently during embryogenesis. In shl2-6 and shl2-7 mutants, the meristem was lost during seed development. Only the shl2-8 mutant could survive after germination, but it showed abnormal initiation pattern and morphology of leaves. In strains bearing the weak alleles, the shoot meristem was composed of a small number of indeterminate cells and ultimately converted into leaf primordium. The shl1-3 mutant showed phenotypes similar to those of shl2-8. Thus SHL2 and SHL1 are required for both initiation and maintenance of shoot meristem. In shl2 mutants, there was a positive correlation between the size of the expression domain of OSH1 representing the number of indeterminate cells, the frequency of shoot meristem initiation, and the duration of meristem survival. Thus the shoot meristem will not initiate in an "all-or-nothing" fashion, but is formed in various degrees depending on the strength of the alleles. Double-mutant analyses indicate that SHL2 functions upstream of SHO to establish proper organization of the shoot meristem.

Homeodomain Proteins↗

PubMed2003

Molecular analysis of the LATERAL SUPPRESSOR gene in Arabidopsis reveals a conserved control mechanism for axillary meristem formation.

In seed plants, shoot branching is initiated by the formation of new meristems in the axils of leaves, which subsequently develop into new axes of growth. This study describes the genetic control of axillary meristem formation by the LATERAL SUPPRESSOR (LAS) gene in Arabidopsis thaliana. las mutants show a novel phenotype that is characterized by the inability to form lateral shoots during vegetative development. The analysis shows that axillary meristem formation is differently regulated during different phases of development. During reproductive development, axillary meristems initiate in close proximity to the shoot apical meristem and do not require LAS function. In contrast, during the vegetative phase, axillary meristems initiate at a distance to the SAM and require LAS function. This control mechanism is conserved between the distantly related species tomato and Arabidopsis. Monitoring the patterns of LAS and SHOOT MERISTEMLESS transcript accumulation allowed us to identify early steps in the development of leaf axil identity, which seem to be a prerequisite for axillary meristem initiation. Other regulators of shoot branching, like REVOLUTA and AUXIN RESISTANT 1, act downstream of LAS. The results are discussed in the context of the "detached meristem" and the "de novo formation" concepts of axillary meristem formation.

Arabidopsis↗