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Bud

A bud is a small, undeveloped protuberance that forms on the stem of a vascular plant, arising from meristem tissue and capable of developing into a flower, leaf, shoot, or combination thereof.^[1] These structures are essential for plant growth and reproduction, originating at nodes where leaves attach to the stem, and they contain undifferentiated cells that enable rapid division and differentiation.^[2] Buds are classified by their position, function, and scale arrangement. Terminal (apical) buds occur at the tips of stems and promote primary growth in length, while axillary (lateral) buds form in the leaf axils and can remain dormant or produce branches.^[3] Adventitious buds arise from non-standard locations like roots or wounds, facilitating vegetative propagation in cuttings.^[4] Functionally, buds divide into vegetative types—such as leaf buds, which consist of a short stem with embryonic leaves and are typically pointed and less plump—and reproductive types like flower buds, which are often rounder and develop into blooms.^[4] Mixed buds combine both leaf and flower primordia, common in many temperate species.^[5] In temperate climates, buds often overwinter protected by scales derived from modified leaves, which prevent desiccation and pathogen entry until activation in spring by environmental cues such as chilling and rising temperatures, promoted by hormones like gibberellins.^[2]^[6] Beyond growth, some enlarged buds serve as food storage, as seen in species like cabbage or onions, while their meristematic activity underpins branching patterns that influence plant architecture and adaptability.^[4]

Anatomy and Structure

External Morphology

A plant bud is defined as a compact, undeveloped shoot that serves as the primary growing point on a stem, containing embryonic tissues such as preformed leaves, stems, or flowers enclosed by protective outer layers.^[7] These structures are essential for vegetative or reproductive development and are typically visible as small protuberances on the plant.^[8] The external morphology of a bud includes several key visible features. At the exterior, bud scales—modified, often leathery leaves—provide protection against environmental stresses, varying from tiny and camouflaged in species like winterberry to furry and gray in magnolias.^[9] The apex, or tip, appears as a pointed or rounded summit housing meristematic tissue responsible for growth, while the base forms the attachment point to the stem or leaf axil, sometimes marked by a scar.^[8] These components give buds a distinct, enclosed appearance that differs from expanded shoots. Buds exhibit variations in shape and size depending on the plant species and bud type, ranging from conical or ovoid forms that are pointed, as seen in many rose buds, to clustered arrangements in oaks.^[10]^[7] For instance, beech buds are characteristically long and shiny, while those in lilacs may occur as double structures, and sizes can span from minuscule and inconspicuous to large and showy, influencing their visibility on the plant.^[9]^[7] Positioning of buds on the plant further defines their external placement. Terminal buds are located at the stem tip, often suppressing lateral growth; axillary buds form in the leaf axil along the stem; and adventitious buds arise at atypical sites such as internodes, roots, or trunks.^[8]^[7] These positions contribute to the overall architectural diversity observed in plant shoots.^[9]

Internal Organization

The internal organization of a plant bud consists primarily of the apical meristem, leaf primordia, and developing vascular tissues, which together form a compact, undifferentiated structure poised for future growth. The apical meristem, located at the bud's tip, comprises undifferentiated cells organized into a tunica-corpus model, where the outer tunica layers undergo anticlinal divisions to produce epidermal tissues, and the inner corpus divides in multiple planes to generate ground and vascular tissues.^[11] These meristematic cells are small, densely cytoplasmic, and actively dividing, providing the source for all subsequent bud components. Enclosed within this meristem are leaf primordia, which are embryonic leaves arising as bulges from the meristem flanks in a phyllotactic pattern, typically stacked vertically in longitudinal section to conserve space.^[11] Vascular tissues within the bud derive from procambial strands, featuring protoxylem elements in the inner regions and protophloem in the outer layers, which connect directly to the parent plant's vascular system but remain incompletely differentiated until bud expansion.^[11] Protective layers surround these internal tissues, including bud scales derived from modified leaf primordia that form a tight, overlapping enclosure. These scales are coated with a waxy cuticle that minimizes water loss and prevents desiccation during adverse conditions.^[12] In certain species, additional protection comes from resin secretions or trichomes (hairs) on the scales, which deter herbivores and further reduce evaporation.^[13] At the cellular level, the bud's interior features densely packed parenchyma cells in the ground tissue, lacking extensive intercellular spaces and featuring thin walls to maintain compactness; fully differentiated vascular bundles are absent, with procambial cells instead forming rudimentary strands that elongate and mature post-dormancy.^[11] In dicots such as apple (Malus domestica), the internal organization exemplifies this layered complexity, with the apical meristem centrally positioned and surrounded by multiple leaf primordia arranged in a vertical stack, often numbering several cataphylls (scale-like) followed by foliage primordia.^[14] This stacking optimizes resource allocation within the enclosed space, ensuring orderly development upon activation.^[15]

Development and Formation

Initiation Process

The initiation of buds in plants primarily occurs through the activity of the shoot apical meristem (SAM), which generates lateral buds via organized cell divisions in the axillary positions at the base of leaves. These axillary meristems arise as small groups of undifferentiated cells derived from the peripheral zone of the SAM, establishing new sites of growth that can develop into branches or reproductive structures.^[16]^[17] Hormonal regulation plays a central role in controlling bud initiation, with auxin produced by the shoot tip exerting inhibitory effects through apical dominance, preventing premature lateral bud formation to prioritize main stem elongation. In contrast, cytokinins actively promote the initiation and early development of axillary meristems by stimulating cell division and counteracting auxin's suppressive signals. This auxin-cytokinin balance ensures that bud sites are established only under appropriate developmental cues, as demonstrated in model systems where exogenous cytokinin application induces ectopic bud formation.^[18] Environmental factors such as photoperiod and temperature significantly influence bud site determination, particularly in temperate species where shortening day lengths signal the transition to bud initiation for overwintering. For instance, in trees like poplar, short days induce the formation of terminal and axillary buds by altering gene expression related to growth cessation. Buds typically form during the active growing season, with the first primordia emerging shortly following leaf axil development; in Arabidopsis thaliana, this process is genetically regulated by the SHOOTMERISTEMLESS (STM) gene, a KNOX transcription factor essential for specifying meristem identity during axillary initiation. These primordia represent the initial structured outgrowths that define early bud architecture.^[19]^[20]^[21]

Growth Stages

Following the initiation of the bud meristem, plant buds undergo a series of distinct growth stages characterized by morphological and cellular changes that lead to visible expansion and the emergence of new structures. These stages typically include swelling, scale burst, and elongation, each driven by environmental cues and internal physiological processes.^[22] The initial stage, swelling, involves increased water uptake into bud cells, primarily through the expansion of central vacuoles, which causes the bud to enlarge without significant cell division. This vacuolation process raises turgor pressure, allowing cells to stretch against the cell wall and increase the bud's overall size by up to several times its dormant volume. Concurrently, vascular tissues begin to differentiate, with procambial cells maturing into xylem for water transport and phloem for nutrient distribution, establishing the vascular framework for subsequent shoot development.^[23]^[24]^[25] As swelling progresses, the protective bud scales—modified leaves that shield the developing tissues—begin to separate or burst open in the scale burst stage, exposing the inner primordia to light and air. This rupture is facilitated by the mechanical pressure from expanding internal tissues and enzymatic weakening of scale attachments, marking the transition from protected dormancy to active growth.^[26] The final elongation stage sees the rapid outgrowth of the stem and leaf primordia, where continued cell expansion and division propel the shoot upward, often reaching several inches in length. Vascular differentiation intensifies here, with xylem and phloem elements fully maturing to support the emerging shoot's transport needs. These stages, building on the embryonic meristem formed during initiation, enable the bud to contribute to the plant's seasonal renewal.^[27]^[28] In temperate regions, bud growth is triggered primarily by spring warming temperatures, occurring rapidly over days to weeks as cumulative heat units accumulate. For instance, lilac (Syringa vulgaris) buds swell and expand in direct response to rising spring temperatures, with branches near warm surfaces advancing their growth earlier than those in cooler microclimates.^[29]^[30] Abnormalities during these growth stages can disrupt normal development, such as fasciation, where meristem disruption—often from physical damage, hormonal imbalance, or infection—leads to flattened, ribbon-like growth instead of cylindrical elongation. This condition typically arises if the apical meristem is altered during swelling or early elongation, resulting in fused organs and altered vascular patterning.^[31]^[32]

Classification

Positional Types

Plant buds are classified into positional types based on their location relative to the stem and other plant structures, which influences their structural development and role in overall plant architecture. Terminal buds, also known as apical buds, are situated at the apex or tip of a stem and are primarily responsible for the elongation of the main shoot axis through primary growth.^[4]^[33] This positioning enables the bud to direct resources toward vertical extension, often resulting in a larger size compared to other buds and exerting dominance over subordinate structures, thereby shaping the plant's primary form.^[8] In conifers, such as pines, terminal buds are prominently located at twig ends and drive the annual extension of branches, contributing to the characteristic whorled structure.^[34] Axillary buds, alternatively termed lateral buds, develop in the axils where leaves attach to the stem, positioning them along the sides of the stem to facilitate potential branching.^[4] This location allows multiple buds per node in certain species, such as in some herbaceous plants, enabling diversified lateral growth when activated.^[4] Axillary buds typically remain dormant under the influence of the terminal bud but can initiate development following removal of apical dominance, as occurs through pruning, leading to increased branching and a bushier habit.^[35] Their dormancy, further explored in plant physiology sections, underscores how positional constraints regulate developmental timing and prevent excessive lateral proliferation during primary shoot extension.^[35] Adventitious buds emerge from atypical locations outside the standard terminal or axillary positions, such as on roots, internodes, or wounded tissues, providing flexibility for regeneration and propagation.^[36] This irregular positioning arises from callus tissue or modified organs, allowing the plant to form new shoots in response to environmental cues or damage, which is crucial for vegetative spread.^[7] In raspberries, adventitious buds develop on roots to produce suckers, enabling clonal expansion through underground rhizomes.^[37] These buds' non-standard sites highlight their structural adaptability, often leading to decentralized growth patterns distinct from the centralized extension of terminal types.^[7]

Functional Types

Plant buds are differentiated by their functional roles in development, primarily categorized as vegetative, reproductive, or mixed, each contributing to specific aspects of growth and reproduction in plants.^[7] Vegetative buds develop into shoots bearing leaves and stems, supporting non-reproductive growth and canopy expansion. These buds are prevalent in woody plants, where they dominate axillary and terminal positions to promote branching and photosynthesis.^[4]^[38] Reproductive buds, often termed flower buds, give rise to inflorescences or individual flowers, facilitating pollination and seed production. They are characterized by internal floral primordia enclosed by protective scales, distinguishing them from vegetative buds by their plumper, more rounded shape in many species.^[38]^[7] Mixed buds integrate both vegetative and reproductive components, yielding shoots with leaves subtending flowers upon expansion. Common in temperate fruit trees such as apples and pears, these buds form seasonally on short spurs and enable synchronized vegetative support for reproductive structures.^[35]

Physiology and Function

Dormancy and Activation

Bud dormancy in plants is categorized into three primary types based on the sources of inhibition: paradormancy, endodormancy, and ecodormancy. Paradormancy involves external inhibition of bud growth by signals from other plant parts, such as auxin produced by apical or elongating shoots that suppresses lateral bud outgrowth.^[39] Endodormancy represents an internal physiological state where buds are inhibited by endogenous factors within the bud itself, independent of external environmental cues.^[40] Ecodormancy occurs when buds are physiologically capable of growth but are restrained by unfavorable external conditions, such as low temperatures or drought.^[40] Breaking dormancy, particularly endodormancy, often requires vernalization through prolonged cold exposure, known as chilling accumulation. For many deciduous fruit trees, this typically involves 400–1200 hours of temperatures below 7°C to fulfill chilling requirements and release internal inhibition.^[41] Gibberellins play a key role in promoting bud activation by stimulating cell elongation and counteracting inhibitory hormones during the transition from dormancy to growth.^[42] At the molecular level, dormancy maintenance and release involve hormonal regulation, including the downregulation of abscisic acid (ABA), which accumulates during endodormancy to enforce quiescence.^[43] Recent advances have identified genes such as EARLY BUD BREAK (EBB) and SHORT VEGETATIVE PHASE 4 (SVP4) that integrate ABA signaling and promote bud break in species like grapevines (Vitis vinifera).^[44] Positional effects, such as those in apical versus lateral buds, can influence dormancy depth, with terminal buds often entering deeper endodormancy.^[45] Incomplete fulfillment of dormancy requirements, such as insufficient chilling due to warmer winters, can lead to premature bud break, exposing tender tissues to late spring frosts and causing significant damage to emerging shoots and flowers.^[46]

Role in Plant Growth

Buds play a central role in determining plant architecture through their regulation of branching patterns, where axillary bud outgrowth synchronizes with leaf and shoot development.^[47] In woody plants like sugar maple, heterophyllous shoots exhibit delayed terminal bud formation but maintain shorter plastochrons, allowing synchronized expansion of leaves and internodes that aligns with environmental gradients such as light availability.^[48] This coordination ensures that bud-initiated modules—repeating units of stem, leaves, and buds—form cohesive shoots, enhancing photosynthetic efficiency and structural stability.^[49] A key mechanism governing bud activity is apical dominance, mediated by correlative inhibition from terminal buds, which suppress lateral bud outgrowth via auxin transport from the shoot apex, thereby shaping overall plant form.^[18] This inhibition promotes monopodial growth, where a dominant central axis elongates continuously from the terminal bud while laterals remain subordinate, as seen in upright trees like beech, fostering height attainment for light competition.^[50] In contrast, reduced dominance enables sympodial growth, characterized by sequential activation of lateral buds after terminal cessation, resulting in branched, spreading architectures in species like cotton fruiting branches, which balance reproduction and vegetative expansion.^[50] Such patterns, first demonstrated by auxin application mimicking terminal bud effects, underscore how buds integrate hormonal signals to control branching density and angle.^[51] Buds further facilitate adaptive growth cycles, enabling determinate patterns in annuals—where shoots complete development within one season via finite bud activation—and indeterminate patterns in perennials, allowing perpetual meristem activity for multi-year persistence.^[52] In perennials, seasonal bud dormancy and reactivation, such as spring bursting in temperate trees, sustain ongoing shoot production, contrasting with annuals' terminal bud exhaustion post-reproduction.^[48] Evolutionarily, this modular bud-based growth confers redundancy, as dormant axillary and below-ground buds regenerate shoots after disturbance, buffering against herbivory or fire and promoting clonal persistence in variable habitats like prairies.^[53] By enabling iterative module production, buds enhance survival through architectural plasticity, a trait amplified in perennials for long-term resource exploitation.^[54]

Applications and Significance

In Horticulture

In horticulture, budding techniques such as T-budding and chip budding serve as efficient methods for grafting, involving the insertion of a single bud from a scion into a prepared slit on the rootstock bark to propagate desirable varieties.^[55]^[56] T-budding, performed during active bark growth, entails making a vertical incision and horizontal cut to form a "T" shape on the rootstock, lifting the bark flaps to nestle the bud shield containing the vascular cambium.^[57] Chip budding, suitable for dormant or growing seasons, removes a thin chip of wood with the bud from the scion and matches it to a similarly shaped cut on the rootstock, allowing quick union formation.^[58] In citrus production, these approaches facilitate the propagation of trees onto rootstocks like trifoliate orange, imparting resistance to diseases such as citrus tristeza virus and Phytophthora root rot, thereby improving orchard longevity and productivity.^[59] Pruning terminal buds disrupts apical dominance mediated by auxins, prompting the activation of axillary buds and fostering denser branching that supports greater fruit set in pome fruits. In apple orchards, this practice—often applied as heading cuts during winter or summer—enhances light penetration and air circulation, reducing disease incidence while boosting flower bud initiation on lateral shoots.^[60] For example, selective removal of dominant terminals in varieties like Honeycrisp promotes balanced canopy development, ensuring consistent fruit quality and size across the tree.^[61] Bud selection in breeding leverages spontaneous mutations known as bud sports to generate novel cultivars with superior traits, bypassing lengthy cross-pollination cycles.^[62] These chimeric mutations, identifiable by distinct leaf or fruit characteristics, are propagated vegetatively to fix the trait; a prominent case is the Red Delicious apple, which arose as a red-pigmented bud sport on a Delicious tree in Iowa in 1917, enabling its widespread adoption for vibrant market appeal and contributing to over 50% of U.S. apple production by the mid-20th century. Such selections have accelerated varietal improvement in horticulture, with vegetative buds commonly targeted for isolating sports in woody perennials.^[63] Bud abortion, triggered by abiotic stresses like water deficit or high temperatures, poses a significant challenge in cultivation by curtailing reproductive and vegetative development, potentially halving potential yields in sensitive crops.^[64] Management strategies include foliar applications of cytokinins, such as benzyladenine, which enhance sink strength in buds, delay senescence, and reduce abortion rates under stress conditions. These treatments activate genes involved in cell division and nutrient mobilization, supporting bud survival without compromising overall plant architecture.^[64]

Ecological Importance

Plant buds play a pivotal role in ecosystem resilience by facilitating vegetative regeneration and population persistence in response to disturbances such as fire, herbivory, and land-use changes. Below-ground bud banks, consisting of dormant meristems on roots, rhizomes, and other subterranean structures, serve as a critical reservoir for regrowth, enabling plants to resprout rapidly after above-ground biomass loss. This regenerative capacity is particularly vital in fire-prone habitats, where bud banks support the resprouter life history strategy, allowing species to recolonize and stabilize vegetation cover post-disturbance.^[53] In grassland and savanna ecosystems, bud banks contribute to community dynamics by buffering against grazing pressure and promoting clonal propagation, which enhances genetic continuity and spatial spread of plant populations. For example, in temperate perennial plants, renewal buds drive spring regrowth and seasonal shoot cycles, maintaining productivity and habitat structure amid fluctuating environmental conditions. The size and composition of bud banks influence invasion resistance, as species with robust bud reserves can outcompete non-resprouting invaders following perturbations.^[65]^[66]^[67] Buds also mediate plant responses to climate variability, with axillary and apical buds regulating branching patterns that optimize light capture and resource allocation in competitive environments. Apical dominance, exerted by terminal buds via hormonal signals, shapes plant architecture to favor height growth in resource-limited settings, thereby influencing interspecific competition and understory diversity in forests and shrublands. Disruptions to bud formation or dormancy, such as those induced by drought or elevated temperatures, can diminish bud bank viability, leading to reduced ecosystem resilience and potential shifts in species composition.^[68]^[69]^[67] Reproductive buds, which develop into flowers, indirectly bolster ecological interactions by supporting pollinator networks and seed dispersal, essential for maintaining biodiversity in angiosperm-dominated communities. Overall, the ecological significance of buds lies in their dual function as sites of growth potential and adaptation, underpinning vegetation recovery and trophic linkages across terrestrial ecosystems.^[66]

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Improving citrus bud grafting efficiency - PMC - NIH
Oct 18, 2023 · Cytokinins stimulate the formation of keikis and one method of propagation is to use keiki paste, a lanolin paste containing a cytokinin, often ...<|control11|><|separator|>
[60]
Pruning Apples and Pears in Home Fruit Plantings
Apr 17, 2023 · Thinning-out cutting (removing an entire limb or shoot) is associated with increased apple flower bud production. Heading-back cutting ( ...
[61]
Training and pruning your home orchard | OSU Extension Service
Pruning young trees can improve structural strength and induce branching. Pruning mature trees can increase their production and improve fruit quality.
[62]
Whole-genome resequencing and transcriptome analyses of four ...
Nov 30, 2023 · 'Red Delicious' is considered to be a special plant material of bud sport, whereas the genetic basis of plant mutants is still unknown.
[63]
Regulation of Flowering Timing by ABA-NnSnRK1 Signaling ...
Apr 10, 2021 · ABA Plays an Important Role in Regulating Lotus Flower Bud Abortion. Phytohormones are thought to play important roles in the PCD process [17].
[64]
Exogenous Gibberellic Acid and Cytokinin Effects on Budbreak ...
Oct 29, 2020 · Budbreak increased linearly with GA3, but yield decreased exponentially with GA3 because of dose-dependent flower abortion. In the third ...
[65]
Diversity and functional differentiation of renewal buds in temperate ...
Aug 12, 2024 · Spring regrowth in temperate perennials relies on renewal buds, which form a key component in the shoot growth cycle.
[66]
Bud banks and their role in vegetative regeneration
We review current knowledge about the role of bud banks in vegetative regeneration and the diversity of their modes of functioning.
[67]
The ecology and significance of below-ground bud banks in plants
Environmental change that depletes the bud bank or prohibits its formation likely results in a loss of vegetation resilience and plant species diversity.
[68]
Growing competitive or tolerant? Significance of apical dominance ...
Dec 2, 2017 · When plants were damaged, the apically dominant ssp. campestris showed a positive population growth rate (λ > 1) and often overcompensatory seed ...
[69]
Belowground bud banks and land use change: roles of vegetation ...
Jan 4, 2024 · Belowground bud banks play integral roles in vegetation regeneration and ecological succession of plant communities; however, human-caused ...