Shoot Apical Meristem Research Articles

Toward Transgene-Free Transposon-Mediated Biological Mutagenesis for Plant Breeding.

Genetic diversity is a key factor for plant breeding. The birth of novel genic and genomic variants is also crucial for plant adaptation in nature. Therefore, the genomes of almost all living organisms possess natural mutagenic mechanisms. Transposable elements (TEs) are a major mutagenic force driving genetic diversity in wild plants and modern crops. The relatively rare TE transposition activity during the thousand-year crop domestication process has led to the phenotypic diversity of many cultivated species. The utilization of TE mutagenesis by artificial and transient acceleration of their activity in a controlled mode is an attractive foundation for a novel type of mutagenesis called TE-mediated biological mutagenesis. Here, I focus on TEs as mutagenic sources for plant breeding and discuss existing and emerging transgene-free approaches for TE activation in plants. Furthermore, I also review the non-randomness of TE insertions in a plant genome and the molecular and epigenetic factors involved in shaping TE insertion preferences. Additionally, I discuss the molecular mechanisms that prevent TE transpositions in germline plant cells (e.g., meiocytes, pollen, egg and embryo cells, and shoot apical meristem), thereby reducing the chances of TE insertion inheritance. Knowledge of these mechanisms can expand the TE activation toolbox using novel gene targeting approaches. Finally, the challenges and future perspectives of plant populations with induced novel TE insertions (iTE plant collections) are discussed.

Local auxin biosynthesis promotes shoot patterning and stem cell differentiation in Arabidopsis shoot apex.

The shoot apical meristem (SAM) of higher plants comprises distinct functional zones. The central zone (CZ) is located at the meristem summit and harbors pluripotent stem cells. Stem cells undergo cell division within the CZ and give rise to descendants, which enter the peripheral zone (PZ) and become recruited into lateral organs. Stem cell daughters that are pushed underneath the CZ form rib meristem (RM). To unravel the mechanism of meristem development, it is essential to know how stem cells adopt distinct cell fates in the SAM. Here, we show that meristem patterning and floral organ primordia formation, besides auxin transport, are regulated by auxin biosynthesis mediated by two closely related genes of the TRYPTOPHAN AMINOTRANSFERASE family. In Arabidopsis SAM, TAA1 and TAR2 played a role in maintaining auxin responses and the identity of PZ cell types. In the absence of auxin biosynthesis and transport, the expression pattern of the marker genes linked to the patterning of the SAM is perturbed. Our results prove that local auxin biosynthesis, in concert with transport, controls the patterning of the SAM into the CZ, PZ and RM.

Two modes of gene regulation by TFL1 mediate its dual function in flowering time and shoot determinacy of Arabidopsis.

Plant organ primordia develop successively at the shoot apical meristem (SAM). In Arabidopsis, primordia formed early in development differentiate into vegetative leaves, whereas those formed later generate inflorescence branches and flowers. TERMINAL FLOWER 1 (TFL1), a negative regulator of transcription, acts in the SAM to delay flowering and to maintain inflorescence meristem indeterminacy. We used confocal microscopy, time-resolved transcript profiling and reverse genetics to elucidate this dual role of TFL1. We found that TFL1 accumulates dynamically in the SAM reflecting its dual function. Moreover, TFL1 represses two major sets of genes. One set includes genes that promote flowering, expression of which increases earlier in tfl1 mutants. The other set is spatially misexpressed in tfl1 inflorescence meristems. The misexpression of these two gene sets in tfl1 mutants depends upon FD transcription factor, with which TFL1 interacts. Furthermore, the MADS-box gene SEPALLATA 4, which is upregulated in tfl1, contributes both to the floral transition and shoot determinacy defects of tfl1 mutants. Thus, we delineate the dual function of TFL1 in shoot development in terms of its dynamic spatial distribution and different modes of gene repression.

A Grafting Procedure for Gossypium hirsutum Based on In Vitro Grown Seedlings

Intervarietal grafting, and in some cases interspecific grafting, practiced mainly in horticultural crops has many practical applications. Grafting can also be a useful tool to perform basic physiological, biochemical, and molecular studies to better understand communication between shoot and root in a row crop, such as cotton. We have established a grafting protocol for upland cotton (Gossypium hirsutum L.) genotypes. Although less efficient, it can be used to create grafts between upland and Pima (G. barbadense) or Old-world cottons (G. arboreum or G. herbaceum). The procedure used 8-day-old seedlings grown in culture under sterile conditions to obtain the scion and rootstock. The scion consisted of the shoot apical meristem with a few leaf primordia, epicotyl, and a small portion of the hypocotyl with the cotyledons removed. The rootstock retained 2 to 3 cm of basal hypocotyl portion with the roots trimmed. Cleft grafts were assembled under aseptic conditions followed by their culture in vitro. Once established, as indicated by the healthy growth of three to four true leaves and roots, the grafts were moved to soil in small pots. After hardening the grafts on a laboratory bench for few weeks, they were transferred to larger pots and grown to maturity in a greenhouse to obtain seeds. The success rate using our protocol ranged from 90 to 100% for Coker 312 genotype and its derivative transgenic and gene edited lines. The method was also applicable to generate intervarietal and interspecific graft unions among various types of cotton plants.

AmPHO2 gene expression and interaction protein screening of snapdragon inoculated with AM fungi under low phosphorus stress

Phosphorus (P) is involved in flowering and fruiting in plants. P deficiency inhibits the formation of plant reproductive organs and affects the process of flower bud differentiation, leads to delayed flowering and affects the number and quality of flowers. Our previous experiment showed that arbuscular mycorrhizal (AM) fungal inoculation could promote snapdragon (Antirrhinum majus L.) uptaking P, prolong the flowering period, and increase the number of flower buds and flowers at the full bloom stage under low P stress. The aim of this study is to preliminarily explore the molecular mechanism of AM fungus enhancing P absorption by snapdragon under low P stress. Four treatments were set up in this experiment, namely, low P level (0.1 mmol /L P), low P level (0.1 mmol /L P) +AM fungi, normal P level (0.7 mmol /L P), and normal P level (0.7 mmol/L P) + AM fungi, to evaluate regulation of AM fungi on the phosphate signaling factor PHOSPHATE 2 (PHO2) and the snapdragon flowering. Results showed that the P content in the roots, shoot apical meristem, buds and flowers of snapdragon inoculated with AM fungi increased and the expression of AmPHO2 decreased, suggesting that mycorrhizal symbiosis may inhibit the expression of AmPHO2. Total 28 AmPHO2 interacting proteins were screened by Yeast two-hybrid technique, among which the thioredoxin AmTrx was involved in the flowering process of the plant, and point-to-point validation and dual luciferase assay of AmTrx was further verified. This study showed that inoculation with AM fungi could negatively regulate AmPHO2, promote P uptake, flowering, and improve flower quality of snapdragon under low P stress.

Regulating bulb dormancy release and flowering in lily through chemical modulation of intercellular communication

Lily is a bulbous plant with an endogenous dormancy trait. Fine-tuning bulb dormancy release is still a challenge in the development of bulb storage technology. In this study, we identified three regulators of symplastic transport, 2,3-Butanedione oxime (BDM), N-Ethyl maleimide (NEM), and 2-Deoxy-D-glucose (DDG), that also regulate bulb dormancy release. We demonstrated that BDM and DDG inhibited callose synthesis between cells and promoted symplastic transport and soluble sugars in the shoot apical meristem (SAM), eventually accelerating bulb dormancy release and flowering in lilies. Conversely, NEM had the opposite effect. These three regulators can be flexibly applied to either accelerate or delay lily bulb dormancy release.

Genome-wide identification of the CclKNOX gene family and functional characterization of CclKNOX3 and CclKNOX5 in citrus shoot development

KNOTTED1-LIKE HOMEOBOX (KNOX) genes are a relatively conserved gene family in plants, and have important regulatory functions in the morphological development of plants. In this study, a total of 9 KNOX proteins were found from citrus and divided into two classes, Class I and Class II. Subsequently, their gene structures, protein characteristics and sequence features were analyzed. Spatial expression analysis showed that most of the genes exhibited a constitutive expression pattern in different tissues of Citrus clementina. Interestingly, the expression pattern of CclKNOX3 was different from other Class I members, as it was mainly expressed in stems, lateral buds and shoot tips. Overexpression of CclKNOX3 led to strong differentiation of shoot apical meristem and curled leaves in transgenic tobacco and lemon. In addition, CclKNOX3 was found to interact with CclKNOX5, localized in the nucleus and have transcriptional inhibitory activity. Further transgenic analysis found that 35S:CclKNOX5 transgenic tobacco showed similar phenotype with 35S:CclKNOX3. Furthermore, yeast one hybrid and luciferase assay confirmed that CclKNOX3 and CclKNOX5 were regulated by CclKNOX4. This study expands our knowledge about the citrus KNOX family and provides valuable informations for future studies on shoot developmental regulation.

Control of stem cell behavior by CLE-JINGASA signaling in the shoot apical meristem in Marchantia polymorpha

Land plants undergo indeterminate growth by the activity of meristems in both gametophyte (haploid) and sporophyte (diploid) generations. In the sporophyte of the flowering plant Arabidopsis thaliana, the apical meristems are located at the shoot and root tips in which a number of regulatory gene homologs are shared for their development, implying deep evolutionary origins. However, little is known about their functional conservation with gametophytic meristems in distantly related land plants such as bryophytes, even thoughgenomic studies have revealed that the subfamily-level diversity of regulatory genes is mostly conserved throughout land plants. Here, we show that a NAM/ATAF/CUC (NAC) domain transcription factor, JINGASA (MpJIN), acts downstream of CLAVATA3 (CLV3)/ESR-related (CLE) peptide signaling and controls stem cell behavior in the gametophytic shoot apical meristem of the liverwort Marchantia polymorpha. In the meristem, strong MpJIN expression was associated with the periclinal cell division at the periphery of the stem cell zone (SCZ), whereas faint MpJIN expression was found at the center of the SCZ. Time course observation indicates that the MpJIN-negative cells are lost from the SCZ and respecified de novo at two separate positions during the dichotomous branching event. Consistently, the induction of MpJIN results in ectopic periclinal cell division in the SCZ and meristem termination. Based on the comparative expression data, we speculate that the function of JIN/FEZ subfamily genes was shared among the shoot apical meristems in the gametophyte and sporophyte generations in early land plants but was lost in certain lineages, including the flowering plant A.thaliana.

CRISPR/Cas9-mediated mutations of FANTASTIC FOUR gene family for creating early flowering mutants in tomato.

Flowering time is of great agricultural importance and the timing and extent of flowering usually determines yield and availability of flowers, fruits and seeds. Identification of genes determining flowering has important practical applications for tomato breeding. Here we demonstrate the roles of the FANTASTIC FOUR (FAF) gene family in regulating tomato flowering time. In this plant-specific gene family, SlFAF1/2a shows a constitutive expression pattern during the transition of the shoot apical meristem (SAM) from vegetative to reproductive growth and significantly influences flowering time. Overexpressing SlFAF1/2a causes earlier flowering compared with the transformations of other genes in the FAF family. SlFAF1/2c also positively regulates tomato flowering, although to a lesser extent. The other members of the SlFAF gene family, SlFAF1/2b, SlFAF3/4a and SlFAF3/4b, are negative regulators of tomato flowering and faf1/2b, faf3/4a and faf3/4b single mutants all display early flowering. We generated a series of early flowering mutants using the CRISPR/Cas9 editing system, and the faf1/2b faf3/4a faf3/4b triple mutant flowering earliest compared with other mutants. More importantly, these mutants show no adverse effect on yield. Our results have uncovered the role of the FAF gene family in regulating tomato flowering time and generated early flowering germplasms for molecular breeding.

The transcription factor HSFA7b controls thermomemory at the shoot apical meristem by regulating ethylene biosynthesis and signaling in Arabidopsis

The shoot apical meristem (SAM) is responsible for overall shoot growth by generating all aboveground structures. Recent research has revealed that the SAM displays an autonomous heat stress (HS) memory of a previous non-lethal HS event. Considering the importance of the SAM for plant growth, it is essential to determine how its thermomemory is mechanistically controlled. Here, we report that HEAT SHOCK TRANSCRIPTION FACTOR A7b (HSFA7b) plays a crucial role in this process in Arabidopsis, as the absence of functional HSFA7b results in the temporal suppression of SAM activity after thermopriming. We found that HSFA7b directly regulates ethylene response at the SAM by binding to the promoter of the key ethylene signaling gene ETHYLENE-INSENSITIVE 3 to establish thermotolerance. Moreover, we demonstrated that HSFA7b regulates the expression of ETHYLENE OVERPRODUCER 1 (ETO1) and ETO1-LIKE 1, both of which encode ethylene biosynthesis repressors, thereby ensuring ethylene homeostasis at the SAM. Taken together, these results reveal a crucial and tissue-specific role for HSFA7b in thermomemory at the Arabidopsis SAM.

Vacuolar Sugar Transporter TMT2 Plays Crucial Roles in Germination and Seedling Development in Arabidopsis

Vacuolar sugar transporters transport sugar across the tonoplast, are major players in maintaining sugar homeostasis, and therefore play vital roles in plant growth, development, and biomass yield. In this study, we analyzed the physiological roles of the tonoplast monosaccharide transporter 2 (TMT2) in Arabidopsis. In contrast to the wild type (WT) that produced uniform seedlings, the tmt2 mutant produced three types of offspring: un-germinated seeds (UnG), seedlings that cannot form true leaves (tmt2-S), and seedlings that develop normally (tmt2-L). Sucrose, glucose, and fructose can substantially, but not completely, rescue the abnormal phenotypes of the tmt2 mutant. Abnormal cotyledon development, arrested true leaf development, and abnormal development of shoot apical meristem (SAM) were observed in tmt2-S seedlings. Cotyledons from the WT and tmt2-L seedlings restored the growth of tmt2-S seedlings through micrografting. Moreover, exogenous sugar sustained normal growth of tmt2-S seedlings with cotyledon removed. Finally, we found that the TMT2 deficiency resulted in growth defects, most likely via changing auxin signaling, target of rapamycin (TOR) pathways, and cellular nutrients. This study unveiled the essential functions of TMT2 for seed germination and initial seedling development, ensuring cotyledon function and mobilizing sugars from cotyledons to seedlings. It also expanded the current knowledge on sugar metabolism and signaling. These findings have fundamental implications for enhancing plant biomass production or seed yield in future agriculture.

Functional Modules in the Meristems: "Tinkering" in Action.

A feature of higher plants is the modular principle of body organisation. One of these conservative morphological modules that regulate plant growth, histogenesis and organogenesis is meristems-structures that contain pools of stem cells and are generally organised according to a common principle. Basic content: The development of meristems is under the regulation of molecular modules that contain conservative interacting components and modulate the expression of target genes depending on the developmental context. In this review, we focus on two molecular modules that act in different types of meristems. The WOX-CLAVATA module, which includes the peptide ligand, its receptor and the target transcription factor, is responsible for the formation and control of the activity of all meristem types studied, but it has its own peculiarities in different meristems. Another regulatory module is the so-called florigen-activated complex, which is responsible for the phase transition in the shoot vegetative meristem (e.g., from the vegetative shoot apical meristem to the inflorescence meristem). The review considers the composition and functions of these two functional modules in different developmental programmes, as well as their appearance, evolution and use in plant breeding.

The Arabidopsis SHORTROOT network coordinates shoot apical meristem development with auxin-dependent lateral organ initiation.

Plants produce new organs post-embryonically throughout their entire life cycle. This is due to stem cells present in the shoot and root apical meristems, the SAM and RAM, respectively. In the SAM, stem cells are located in the central zone where they divide slowly. Stem cell daughters are displaced laterally and enter the peripheral zone, where their mitotic activity increases and lateral organ primordia are formed. How the spatial arrangement of these different domains is initiated and controlled during SAM growth and development, and how sites of lateral organ primordia are determined in the peripheral zone is not yet completely understood. We found that the SHORTROOT (SHR) transcription factor together with its target transcription factors SCARECROW (SCR), SCARECROW-LIKE23 (SCL23) and JACKDAW (JKD), promotes formation of lateral organs and controls shoot meristem size. SHR, SCR, SCL23, and JKD are expressed in distinct, but partially overlapping patterns in the SAM. They can physically interact and activate expression of key cell cycle regulators such as CYCLIND6;1 (CYCD6;1) to promote the formation of new cell layers. In the peripheral zone, auxin accumulates at sites of lateral organ primordia initiation and activates SHR expression via the auxin response factor MONOPTEROS (MP) and auxin response elements in the SHR promoter. In the central zone, the SHR-target SCL23 physically interacts with the key stem cell regulator WUSCHEL (WUS) to promote stem cell fate. Both SCL23 and WUS expression are subject to negative feedback regulation from stem cells through the CLAVATA signaling pathway. Together, our findings illustrate how SHR-dependent transcription factor complexes act in different domains of the shoot meristem to mediate cell division and auxin dependent organ initiation in the peripheral zone, and coordinate this activity with stem cell maintenance in the central zone of the SAM.

Metabolome and transcriptome analyses identify the underground rhizome growth through the regulation of rhizome apices in Panax ginseng

Panax ginseng C. A. Mey. is a well-known oriental medicine as a stimulant and dietary supplement. The ginseng rhizome, as one of the principal medicinal organs, plays an important role in aboveground organ growth and root development, although no studies on ginseng rhizome growth have been recorded. In this study, we identified that shoot apical meristem (SAM) was included in the rhizome bud, which could regulate the underground rhizome growth of ginseng. In microscopic examination, ginseng SAM included the central zone, peripheral zone, rib zone, and organizing center; secondary root structures were observed in ginseng main roots and lateral roots; primary root structures were observed in ginseng fibrous roots; rhizome had a stem structure with pith and discontinuous cambium. The metabolites in different underground ginseng organs were identified using ultra-performance liquid chromatography-electrospray ionization-tandem mass spectrometry (UPLC-ESI-MS/MS), with oleanane (OA)-type ginsenosides accumulating primarily in the rhizome buds. In addition, ginseng rhizome buds contained a low level of 3-indoleacetonitrile (IAN, one type of auxin), a high amount of abscisic acid (ABA), and a high level of jasmonoyl-isoleucine (JA-Ile, active form of jasmonic acid). ABA and JA-Ile could drastically limit the shooting rate of ginseng embryonic calli, however IAN boosted the rooting rate. Furthermore, the expression of genes involved in ginsenoside biosynthesis, phytohormone production, and shoot stem cell regulatory pathways were studied. This is the first study of SAM in medicinal herbs, and we believe that the results may offer new information about the development of ginseng underground rhizomes.

A new mathematical model of phyllotaxis to solve the genuine puzzle spiromonostichy.

Arrangement of plant leaves around the stem, termed phyllotaxis, exhibits beautiful and mysterious regularities and has been one of the most attractive subjects of biological pattern formation. After the long history of studies on phyllotaxis, it is now widely accepted that the inhibitory effect of existing leaf primordia on new primordium formation determines phyllotactic patterning. However, costoid phyllotaxis unique to Costaceae of Zingiberales, displaying spiromonostichy characterized by a steep spiral with a small divergence angle, seems to disagree with the inhibitory effect-based mechanism and has remained as a "genuine puzzle". We developed a new mathematical model, hypothesizing that each leaf primordium exerts not only the inhibitory effect but also some inductive effect. Computer simulations with the new model successfully generated a spiromonostichous pattern when these two effects met a certain relationship. The obtained spiromonostichy matched the real costoid phyllotaxis observed with Costus megalobractea, particularly for the decrease of the divergence angle associated with the enlargement of the shoot apical meristem. The new model was also shown to be able to produce a one-sided distichous pattern that is seen in phyllotaxis of a few plants of Zingiberales and has never been addressed in the previous model studies. These results implicated inductive as well as inhibitory mechanisms in phyllotactic patterning, at least in Zingiberales.

Salicylic acid and RNA interference mediate antiviral immunity of plant stem cells

Stem cells are essential for the development and organ regeneration of multicellular organisms, so their infection by pathogenic viruses must be prevented. Accordingly, mammalian stem cells are highly resistant to viral infection due to dedicated antiviral pathways including RNA interference (RNAi). In plants, a small group of stem cells harbored within the shoot apical meristem generate all postembryonic above-ground tissues, including the germline cells. Many viruses do not proliferate in these cells, yet the molecular bases of this exclusion remain only partially understood. Here, we show that a plant-encoded RNA-dependent RNA polymerase, after activation by the plant hormone salicylic acid, amplifies antiviral RNAi in infected tissues. This provides stem cells with RNA-based virus sequence information, which prevents virus proliferation. Furthermore, we find RNAi to be necessary for stem cell exclusion of several unrelated RNA viruses, despite their ability to efficiently suppress RNAi in the rest of the plant. This work elucidates a molecular pathway of great biological and economic relevance and lays the foundations for our future understanding of the unique systems underlying stem cell immunity.

A quantitative morphospace of multicellular organ design in the plant Arabidopsis

Organ function emerges from the interactions between their constituent cells. The investigation of cellular organization can provide insight into organ function following structure-function relationships. Here, we investigate the extent to which properties in cellular organization can arise "for free" as an emergent property of embedding cells in space versus those that are actively generated by patterning processes. Default cellular configurations were established using three-dimensional (3D) digital tissue models. Network-based analysis of these synthetic cellular assemblies established a quantitative topological baseline of cellular organization, granted by virtue of passive spatial packing and the minimal amount of order that emerges for free in tessellated tissues. A 3D cellular-resolution digital tissue atlas for the model plant species Arabidopsis was generated, and the extent to which the organs in this organism conform to the default configurations was established through statistical comparisons with digital tissue models. Cells in different tissues of Arabidopsis do not conform to random packing arrangements to varying degrees. Most closely matching the random models was the undifferentiated shoot apical meristem (SAM) from which aerial organs emanate. By contrast, leaf and sepal tissue showed the greatest deviation from this baseline, suggesting these to be the most "complex" tissues in Arabidopsis. Investigation of the patterning principles responsible for the gap between these tissues and default patterns revealed cell elongation and the introduction of air spaces to contribute toward additional organ patterning complexity. This work establishes a quantitative morphospace to understand the principles of organ construction and its diversity within a single organism.

Regulation of Flowering Time and Other Developmental Plasticities by 3' Splicing Factor-Mediated Alternative Splicing in Arabidopsis thaliana.

Plants, as sessile organisms, show a high degree of plasticity in their growth and development and have various strategies to cope with these alterations under continuously changing environments and unfavorable stress conditions. In particular, the floral transition from the vegetative and reproductive phases in the shoot apical meristem (SAM) is one of the most important developmental changes in plants. In addition, meristem regions, such as the SAM and root apical meristem (RAM), which continually generate new lateral organs throughout the plant life cycle, are important sites for developmental plasticity. Recent findings have shown that the prevailing type of alternative splicing (AS) in plants is intron retention (IR) unlike in animals; thus, AS is an important regulatory mechanism conferring plasticity for plant growth and development under various environmental conditions. Although eukaryotes exhibit some similarities in the composition and dynamics of their splicing machinery, plants have differences in the 3' splicing characteristics governing AS. Here, we summarize recent findings on the roles of 3' splicing factors and their interacting partners in regulating the flowering time and other developmental plasticities in Arabidopsis thaliana.

Single-cell transcriptomic analysis of pea shoot development and cell-type-specific responses to boron deficiency.

Understanding how nutrient stress impacts plant growth is fundamentally important to the development of approaches to improve crop production under nutrient limitation. Here we applied single-cell RNA sequencing to shoot apices of Pisum sativum grown under boron (B) deficiency. We identified up to 15 cell clusters based on the clustering of gene expression profiles and verified cell identity with cell-type-specific marker gene expression. Different cell types responded differently to B deficiency. Specifically, the expression of photosynthetic genes in mesophyll cells (MCs) was down-regulated by B deficiency, consistent with impaired photosynthetic rate. Furthermore, the down-regulation of stomatal development genes in guard cells, including homologs of MUTE and TOO MANY MOUTHS, correlated with a decrease in stomatal density under B deficiency. We also constructed the developmental trajectory of the shoot apical meristem (SAM) cells and a transcription factor interaction network. The developmental progression of SAM to MC was characterized by up-regulation of genes encoding histones and chromatin assembly and remodeling proteins including homologs of FASCIATA1 (FAS1) and SWITCH DEFECTIVE/SUCROSE NON-FERMENTABLE (SWI/SNF) complex. However, B deficiency suppressed their expression, which helps to explain impaired SAM development under B deficiency. These results represent a major advance over bulk-tissue RNA-seq analysis in which cell-type-specific responses are lost and hence important physiological responses to B deficiency are missed. The reported findings reveal strategies by which plants adapt to B deficiency thus offering breeders a set of specific targets for genetic improvement. The reported approach and resources have potential applications well beyond P. sativum species and could be applied to various legumes to improve their adaptability to multiple nutrient or abiotic stresses.

Stem Cells: Engines of Plant Growth and Development.

The development of both animals and plants relies on populations of pluripotent stem cells that provide the cellular raw materials for organ and tissue formation. Plant stem cell reservoirs are housed at the shoot and root tips in structures called meristems, with the shoot apical meristem (SAM) continuously producing aerial leaf, stem, and flower organs throughout the life cycle. Thus, the SAM acts as the engine of plant development and has unique structural and molecular features that allow it to balance self-renewal with differentiation and act as a constant source of new cells for organogenesis while simultaneously maintaining a stem cell reservoir for future organ formation. Studies have identified key roles for intercellular regulatory networks that establish and maintain meristem activity, including the KNOX transcription factor pathway and the CLV-WUS stem cell feedback loop. In addition, the plant hormones cytokinin and auxin act through their downstream signaling pathways in the SAM to integrate stem cell activity and organ initiation. This review discusses how the various regulatory pathways collectively orchestrate SAM function and touches on how their manipulation can alter stem cell activity to improve crop yield.