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European beech (Fagus sylvatica) bud

In botany, a bud is an undeveloped or embryonic shoot and normally occurs in the axil of a leaf or at the tip of a stem. Once formed, a bud may remain for some time in a dormant condition, or it may form a shoot immediately. Buds may be specialized to develop flowers or short shoots or may have the potential for general shoot development. The term bud is also used in zoology, where it refers to an outgrowth from the body which can develop into a new individual.

Overview

[edit]
Inflorescence bud scales in Halesia carolina

The buds of many woody plants, especially in temperate or cold climates, are protected by a covering of modified leaves called scales which tightly enclose the more delicate parts of the bud. Many bud scales are covered by a gummy substance which serves as added protection. When the bud develops, the scales may enlarge somewhat but usually just drop off, leaving a series of horizontally-elongated scars on the surface of the growing stem. By means of these scars one can determine the age of any young branch, since each year's growth ends in the formation of a bud, the formation of which produces an additional group of bud scale scars. Continued growth of the branch causes these scars to be obliterated after a few years so that the total age of older branches cannot be determined by this means.[citation needed]

In many plants, scales do not form over the bud, and the bud is then called a naked bud.[1] The minute underdeveloped leaves in such buds are often excessively hairy. Naked buds are found in some shrubs, like some species of the Sumac and Viburnums (Viburnum alnifolium and V. lantana)[2] and in herbaceous plants. In many of the latter, buds are even more reduced, often consisting of undifferentiated masses of cells in the axils of leaves. A terminal bud occurs on the end of a stem and lateral buds are found on the side. A head of cabbage (see Brassica) is an exceptionally large terminal bud, while Brussels sprouts are large lateral buds.[citation needed]

Since buds are formed in the axils of leaves, their distribution on the stem is the same as that of leaves. There are alternate, opposite, and whorled buds, as well as the terminal bud at the tip of the stem. In many plants buds appear in unexpected places: these are known as adventitious buds.[3]

Often it is possible to find a bud in a remarkable series of gradations of bud scales. In the buckeye, for example, one may see a complete gradation from the small brown outer scale through larger scales which on unfolding become somewhat green to the inner scales of the bud, which are remarkably leaf-like. Such a series suggests that the scales of the bud are in truth leaves, modified to protect the more delicate parts of the plant during unfavorable periods.[citation needed]

Types of buds

[edit]
Plant buds classification
Terminal, vegetative bud of Ficus carica

Buds are often useful in the identification of plants, especially for woody plants in winter when leaves have fallen.[4] Buds may be classified and described according to different criteria: location, status, morphology, and function.[citation needed]

Botanists commonly use the following terms:

  • for location:[citation needed]
    • terminal, when located at the tip of a stem (apical is equivalent but rather reserved for the one at the top of the plant);
    • axillary, when located in the axil of a leaf (lateral is the equivalent but some adventitious buds may be lateral too);
    • adventitious, when located elsewhere, for example on the trunk or roots (some adventitious buds may be former axillary ones that are reduced and hidden under the bark, while other adventitious buds are completely new formed ones).
  • for status:[citation needed]
    • accessory, for secondary buds formed besides a principal bud (axillary or terminal);
    • resting, for a bud that forms at the end of a growth season, and then lies dormant until the onset of the next growth season;
    • dormant or latent, for buds whose growth has been delayed for a rather long time. The term is usable as a synonym of resting, but is better employed for buds waiting undeveloped for years, for example epicormic buds;
    • pseudoterminal, for an axillary bud taking over the function of a terminal bud (characteristic of species whose growth is sympodial: terminal bud dies and is replaced by the closer axillary bud, for examples beech, persimmon, Platanus have sympodial growth).
  • for morphology:
    • scaly or covered (perulate), when scales, also referred to as a perule (lat. perula, perulaei) (which are in fact transformed and reduced leaves) cover and protect the embryonic parts;
    • naked, when not covered by scales;
    • hairy, when also protected by hairs (it may apply either to scaly or to naked buds).
  • for function:[citation needed]
    • vegetative, only containing vegetative structures: a leaf bud is an embryonic shoot containing leaves;
    • reproductive, only containing embryonic flower(s): a flower bud contains a single flower while an inflorescence bud contains an inflorescence;
    • mixed, containing both embryonic leaves and flower(s).
[edit]

References

[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia

Bud

View on Grokipedia
from Grokipedia
A bud is a small, undeveloped protuberance that forms on the stem of a , arising from tissue and capable of developing into a flower, , shoot, or combination thereof. 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. Buds are classified by their position, function, and scale arrangement. Terminal (apical) buds occur at the tips of stems and promote primary growth in , while axillary (lateral) buds form in the leaf axils and can remain dormant or produce branches. Adventitious buds arise from non-standard locations like or wounds, facilitating vegetative propagation in cuttings. Functionally, buds divide into vegetative types—such as 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. Mixed buds combine both leaf and flower primordia, common in many temperate . In temperate climates, buds often overwinter protected by scales derived from modified leaves, which prevent and entry until activation in spring by environmental cues such as chilling and rising temperatures, promoted by hormones like . Beyond growth, some enlarged buds serve as , as seen in like or onions, while their meristematic activity underpins branching patterns that influence architecture and adaptability.

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. These structures are essential for vegetative or reproductive development and are typically visible as small protuberances on the . 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. 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. 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 buds, to clustered arrangements in oaks. For instance, 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. 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. These positions contribute to the overall architectural diversity observed in plant shoots.

Internal Organization

The internal organization of a plant bud consists primarily of the , leaf primordia, and developing vascular tissues, which together form a compact, undifferentiated structure poised for future growth. The , 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. 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 , typically stacked vertically in longitudinal section to conserve space. 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. 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 that minimizes water loss and prevents during adverse conditions. In certain species, additional protection comes from secretions or trichomes (hairs) on the scales, which deter herbivores and further reduce evaporation. At the cellular level, the bud's interior features densely packed cells in the , 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. In dicots such as apple (Malus domestica), the internal organization exemplifies this layered complexity, with the apical centrally positioned and surrounded by multiple leaf primordia arranged in a vertical stack, often numbering several cataphylls (scale-like) followed by foliage primordia. This stacking optimizes within the enclosed space, ensuring orderly development upon activation.

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. Hormonal regulation plays a central role in controlling bud initiation, with produced by the shoot tip exerting inhibitory effects through , preventing premature lateral bud formation to prioritize main stem elongation. In contrast, s actively promote the initiation and early development of axillary meristems by stimulating 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. Environmental factors such as photoperiod and significantly influence bud site determination, particularly in temperate 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 related to growth cessation. Buds typically form during the active , with the first primordia emerging shortly following axil development; in , this process is genetically regulated by the SHOOTMERISTEMLESS (STM) gene, a KNOX essential for specifying identity during axillary initiation. These primordia represent the initial structured outgrowths that define early bud architecture.

Growth Stages

Following the initiation of the bud , 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. The initial stage, swelling, involves increased uptake into bud cells, primarily through the expansion of central vacuoles, which causes the bud to enlarge without significant . This vacuolation process raises , allowing cells to stretch against the 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 for transport and for nutrient distribution, establishing the vascular framework for subsequent shoot development. 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. 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. 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 () 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. Abnormalities during these growth stages can disrupt normal development, such as , where disruption—often from physical damage, hormonal imbalance, or —leads to flattened, ribbon-like growth instead of cylindrical elongation. This condition typically arises if the apical is altered during swelling or early elongation, resulting in fused organs and altered vascular patterning.

Classification

Positional Types

Plant buds are classified into positional types based on their location relative to the stem and other 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. 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. In , such as pines, terminal buds are prominently located at twig ends and drive the annual extension of branches, contributing to the characteristic whorled structure. 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. This location allows multiple buds per node in certain species, such as in some herbaceous plants, enabling diversified lateral growth when activated. 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. Their dormancy, further explored in plant physiology sections, underscores how positional constraints regulate developmental timing and prevent excessive lateral proliferation during primary shoot extension. Adventitious buds emerge from atypical locations outside the standard terminal or axillary positions, such as on , internodes, or wounded tissues, providing flexibility for regeneration and . This irregular positioning arises from tissue or modified organs, allowing the plant to form new shoots in response to environmental cues or damage, which is crucial for vegetative spread. In raspberries, adventitious buds develop on to produce suckers, enabling clonal expansion through underground rhizomes. These buds' non-standard sites highlight their structural adaptability, often leading to decentralized growth patterns distinct from the centralized extension of terminal types.

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 in . Vegetative buds develop into shoots bearing leaves and stems, supporting non-reproductive growth and canopy expansion. These buds are prevalent in woody , where they dominate axillary and terminal positions to promote branching and . Reproductive buds, often termed flower buds, give rise to inflorescences or individual flowers, facilitating and 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. 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.

Physiology and Function

Dormancy and Activation

Bud in 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 parts, such as produced by apical or elongating shoots that suppresses lateral bud outgrowth. Endodormancy represents an internal physiological state where buds are inhibited by endogenous factors within the bud itself, independent of external environmental cues. Ecodormancy occurs when buds are physiologically capable of growth but are restrained by unfavorable external conditions, such as low temperatures or . Breaking , particularly endodormancy, often requires through prolonged cold exposure, known as chilling accumulation. For many fruit trees, this typically involves 400–1200 hours of temperatures below 7°C to fulfill chilling requirements and release internal inhibition. play a key role in promoting bud by stimulating cell elongation and counteracting inhibitory hormones during the transition from dormancy to growth. At the molecular level, maintenance and release involve hormonal regulation, including the downregulation of (ABA), which accumulates during endodormancy to enforce quiescence. 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 (). Positional effects, such as those in apical versus lateral buds, can influence depth, with terminal buds often entering deeper endodormancy. Incomplete fulfillment of dormancy requirements, such as insufficient chilling due to warmer winters, can lead to premature bud break, exposing tender tissues to frosts and causing significant damage to emerging shoots and flowers.

Role in Plant Growth

Buds play a central role in determining plant through their regulation of branching patterns, where outgrowth synchronizes with and shoot development. In woody 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. This coordination ensures that bud-initiated modules—repeating units of stem, leaves, and buds—form cohesive shoots, enhancing and . 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. 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. 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. Such patterns, first demonstrated by auxin application mimicking terminal bud effects, underscore how buds integrate hormonal signals to control branching density and angle. Buds further facilitate adaptive growth cycles, enabling determinate patterns in annuals—where shoots complete development within one via finite bud —and indeterminate patterns in perennials, allowing perpetual activity for multi-year persistence. In perennials, seasonal bud and reactivation, such as spring bursting in temperate trees, sustain ongoing shoot production, contrasting with annuals' terminal bud exhaustion post-reproduction. Evolutionarily, this modular bud-based growth confers redundancy, as dormant axillary and below-ground buds regenerate shoots after disturbance, buffering against herbivory or and promoting clonal persistence in variable habitats like prairies. By enabling iterative module production, buds enhance through architectural plasticity, a trait amplified in perennials for long-term resource exploitation.

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. 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. 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. 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. Pruning terminal buds disrupts mediated by auxins, prompting the activation of axillary buds and fostering denser branching that supports greater set in 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. For example, selective removal of dominant terminals in varieties like promotes balanced canopy development, ensuring consistent quality and size across the . Bud selection in breeding leverages spontaneous mutations known as bud sports to generate novel cultivars with superior traits, bypassing lengthy cross-pollination cycles. These chimeric , identifiable by distinct or 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 in , 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 , with vegetative buds commonly targeted for isolating sports in woody perennials. 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. Management strategies include foliar applications of cytokinins, such as benzyladenine, which enhance sink strength in buds, delay , and reduce abortion rates under stress conditions. These treatments activate genes involved in and nutrient mobilization, supporting bud survival without compromising overall plant architecture.

Ecological Importance

Plant buds play a pivotal in resilience by facilitating vegetative regeneration and population persistence in response to disturbances such as , herbivory, and land-use changes. Below-ground bud banks, consisting of dormant meristems on , rhizomes, and other subterranean structures, serve as a critical for regrowth, enabling to resprout rapidly after above-ground loss. This regenerative capacity is particularly vital in fire-prone habitats, where bud banks support the resprouter life history , allowing to recolonize and stabilize cover post-disturbance. In and ecosystems, bud banks contribute to community dynamics by buffering against pressure and promoting clonal propagation, which enhances genetic continuity and spatial spread of populations. For example, in temperate plants, renewal buds drive spring regrowth and seasonal shoot cycles, maintaining productivity and structure amid fluctuating environmental conditions. The size and composition of bud banks influence resistance, as with robust bud reserves can outcompete non-resprouting invaders following perturbations. Buds also mediate plant responses to climate variability, with axillary and apical buds regulating branching patterns that optimize light capture and 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 and diversity in forests and shrublands. Disruptions to bud formation or , such as those induced by or elevated temperatures, can diminish bud bank viability, leading to reduced resilience and potential shifts in composition. Reproductive buds, which develop into flowers, indirectly bolster ecological interactions by supporting networks and , essential for maintaining in angiosperm-dominated communities. Overall, the ecological significance of buds lies in their dual function as sites of growth potential and , underpinning vegetation recovery and trophic linkages across terrestrial ecosystems.

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