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Plant stem

The plant stem is the main structural axis of the shoot system in vascular plants, typically extending above ground from the roots and serving as the primary conduit for water, nutrients, and photosynthetic products throughout the plant. It typically consists of nodes—points where leaves, buds, or branches attach—and internodes, the segments between nodes that contribute to the stem's length and rigidity. Anatomically, stems are organized into three primary tissue systems: the dermal tissue (outer epidermis or periderm for protection), the (xylem for water transport and phloem for nutrient distribution), and the (for storage and support). The primary functions of stems include mechanical support for aerial organs like leaves and flowers, transport of essential substances via the vascular system, and in some species, storage of carbohydrates or water. Stems exhibit diverse forms, from herbaceous and flexible in annuals to woody and lignified in trees, adapting to environmental needs such as climbing, protection, or .

Terminology and Basic Features

Key Terms

In botany, a stem is the main axis of a vascular plant's shoot system, typically above ground, that supports leaves, buds, and reproductive structures while facilitating the transport of water, nutrients, and photosynthates between roots and other organs. A node refers to the specific point along the stem where leaves, branches, or buds attach, serving as a key site for lateral growth and branching. The region between two consecutive nodes is termed the internode, which represents the elongated portion of the stem responsible for overall height and spacing of attachments. A bud is an undeveloped shoot or flower enclosed by protective scales, with two primary types: the terminal bud, located at the apex of the stem to promote apical growth and dominance, and the axillary bud, positioned in the leaf axil at a node, enabling lateral branching and potentially suppressed by the terminal bud. A vascular bundle consists of a discrete strand of conducting tissues—primarily xylem for water transport and phloem for nutrient distribution—often arranged in a cylindrical pattern within the stem and sometimes enclosed by a sheath. Lenticels are specialized, porous regions in the periderm of woody stems, composed of loosely packed cells that facilitate gas exchange between the internal tissues and the atmosphere. The vascular cylinder, also known as the stele, denotes the central core of vascular and ground tissues in the stem (and roots), encompassing the vascular bundles and surrounding parenchyma to form a structural and conductive unit. Specialized stem types include the , a horizontal underground stem that produces roots and shoots at its nodes; the term derives from the Ancient Greek rhízōma, meaning "mass of roots," reflecting its root-like appearance despite being a stem. Similarly, a stolon is a slender, above-ground horizontal stem that roots at nodes to form new plants; its name originates from the Latin stolo, denoting a "shoot" or "sucker," a usage adopted in botany from classical descriptions of propagating branches. Terminologically, stems are distinguished from roots by the presence of nodes and internodes, which roots lack, and from leaves by their role as axial supports rather than flattened photosynthetic organs attached at nodes.

External Morphology

The external morphology of plant stems encompasses the observable surface characteristics that facilitate support, protection, and interaction with the environment. The epidermis forms the outermost layer, consisting of a single sheet of cells often coated with a waxy cuticle to minimize water loss and provide a barrier against pathogens. Nodes appear as distinct swellings or joints along the stem where leaves, buds, or branches attach, serving as sites for lateral growth initiation. Internodes constitute the elongated segments between consecutive nodes, varying in length based on species and environmental conditions to optimize light capture or structural stability. The apical meristem, located at the stem tip, drives primary elongation through continuous cell division. In woody stems, the is eventually replaced by , a tough, multilayered protective covering that develops from corky tissues and shields the inner structures from mechanical damage and . Surface variations enhance functionality; for instance, trichomes or hairs cover the epidermis in many to reduce or deter herbivores, while thorns—modified stem projections—provide , as seen in hawthorns. Ridges or grooves often sculpt the stem surface for increased surface area or structural reinforcement, and lenticels manifest as raised, lens-shaped pores in , enabling in woody tissues where stomata are absent. Stem orientation and shape adapt to ecological niches: erect forms, such as those in sunflowers, support upright growth for , while prostrate stems in groundcovers like strawberries spread horizontally for vegetative propagation. Climbing stems, exemplified by twining vines like or pole beans, coil or adhere to supports for elevation. Herbaceous stems remain soft, green, and flexible throughout their lifespan, contrasting with woody exteriors that harden via , forming durable trunks and branches in trees like oaks. These external traits often correlate with underlying vascular bundles, which influence stem rigidity and patterning.

Anatomy of Stems

Primary Structure

The primary structure of a plant stem refers to the initial organization of tissues derived from the shoot apical during primary growth, before any secondary thickening occurs. This embryonic arrangement arises from the differentiation of three primary s: the protoderm, procambium, and ground . The protoderm gives rise to the outermost epidermal layer, providing a protective covering for the young stem. Meanwhile, the procambium develops into the primary vascular tissues, consisting of and organized within vascular bundles, which facilitate water and nutrient transport. The ground differentiates into the internal ground tissues, including the and , which contribute to storage and structural support. In a typical primary stem cross-section, the tissues are arranged in concentric layers from the periphery inward. The epidermis forms a single layer of tightly packed cells, often covered by a cuticle to minimize water loss and protect against pathogens. Beneath it lies the cortex, a region of parenchyma cells that serves for temporary storage of nutrients and starch, as well as providing mechanical support through collenchyma or sclerenchyma in some areas. The vascular bundles, embedded within or at the inner boundary of the cortex, contain primary xylem toward the center and primary phloem on the outer side, separated by cambium in open bundles capable of further division. At the core is the pith, composed of large, thin-walled parenchyma cells that store water, nutrients, and sometimes starch, occupying the central portion of the stem. The arrangement of vascular tissues varies among vascular , reflecting evolutionary adaptations. In dicotyledons, the vascular bundles are organized in a ring around the , forming an eustele that allows for efficient radial transport and potential . In contrast, monocotyledons exhibit an atactostele, where numerous vascular bundles are scattered throughout the , providing distributed support and conduction suited to herbaceous growth. These patterns originate from the procambial strands during stem elongation, ensuring coordinated tissue development from the apical . Differences in bundle arrangement are more pronounced across groups, as detailed in .

Secondary Structure

Secondary growth in plant stems occurs through the activity of lateral meristems, primarily the , which enables radial expansion and the development of woody tissues in many species. The forms a continuous sheath of undifferentiated cells between the primary and , dividing periclinally to produce secondary toward the interior and secondary toward the exterior. This process thickens the stem, with secondary accumulating as wood that provides and water conduction, while secondary facilitates nutrient transport. Additionally, the , or phellogen, arises in the or pericycle and produces the periderm, a protective outer layer consisting of phellem (), phelloderm, and phellogen itself, which replaces the as the stem expands. In temperate climates, secondary xylem formation results in distinct annual rings visible in cross-sections of woody stems, reflecting seasonal variations in growth. Each ring comprises earlywood, formed in spring under favorable conditions with larger, thinner-walled vessels or tracheids for efficient , and latewood, produced in summer with smaller, thicker-walled cells that contribute to density and strength. Ring width varies based on environmental factors such as and , which influence cambial activity, as well as the plant's age, where younger stems typically exhibit wider rings that narrow with maturity due to competition for resources. These rings serve as records of annual growth increments, with narrower rings often indicating stressful conditions like . Secondary xylem tissues differentiate into sapwood and heartwood, with ray cells facilitating radial transport throughout. Sapwood, the outer pale layer, consists of functional, living and conducting elements that actively transport water and minerals longitudinally. In contrast, heartwood forms the inner, darker core where cells become non-conductive, filled with extractives like , gums, and resins that enhance decay resistance and structural integrity. Ray cells, originating from cambial initials, extend radially as parenchyma strands, enabling lateral movement of nutrients, water, and storage compounds between xylem vessels and the cambium. This organization supports the longevity of perennial plants by balancing conduction, storage, and protection.

Comparative Anatomy

Dicotyledonous Stems

Dicotyledonous stems are characterized by a primary structure in which discrete vascular bundles are arranged in a ring around the periphery of the central pith, a condition known as an eustele. Each vascular bundle consists of primary xylem located toward the center and primary phloem toward the periphery, often separated by a layer of procambium that can develop into fascicular cambium. The epidermis forms the outermost protective layer, typically covered by a cuticle, while the cortex beneath it comprises parenchyma cells, often with collenchyma strands providing mechanical support just below the epidermis. The central pith is composed of large, thin-walled parenchyma cells that store nutrients and water. In young herbaceous dicots, such as the sunflower (Helianthus annuus), a cross-section reveals approximately 8 to 15 vascular bundles arranged in a distinct ring, enclosing the and surrounded by the . The in sunflower stems features prominent collenchyma ridges for added strength, contributing to the stem's toughness despite its non-woody nature, with the occupying a significant portion of the stem's interior. An innermost layer of the , known as the starch sheath, contains -laden cells that function in reserve storage, analogous to an in regulating internal transport. Secondary growth in dicotyledonous stems begins when the fascicular within each connects with interfascicular cambium from the rays, forming a continuous cylindrical ring of that encircles the . This lateral produces secondary cells toward the interior and secondary cells toward the exterior, resulting in radial thickening of the . Over time, the extensive accumulation of secondary forms the bulk of the woody tissue, creating durable trunks in perennial dicots and producing annual growth rings from seasonal variations in cell size and density. In woody dicots like the (Quercus spp.), a cross-section of an older displays concentric layers of secondary , with distinct annual rings marking each year's growth: earlywood with large, thin-walled vessels for efficient water conduction in , followed by denser latewood with smaller cells for support in summer./03:_Plant_Structure/3.03:_Stems/3.3.03:_Secondary_Stem) The secondary is confined to a thin layer beneath the cork cambium-derived , while the inner secondary differentiates into heartwood (non-functional, lignified core) and sapwood (active outer zone). This robust enables oaks and similar to achieve substantial girth, supporting tall canopies and long lifespans./03:_Plant_Structure/3.03:_Stems/3.3.03:_Secondary_Stem)

Monocotyledonous Stems

Monocotyledonous stems exhibit a primary structure adapted primarily for herbaceous growth, lacking the vascular cambium that enables secondary thickening in many other plants. The vascular bundles, which conduct water, nutrients, and sugars, are distributed irregularly throughout the stem, a condition known as an atactostele. This scattered arrangement contrasts with the ring-like organization of vascular bundles in dicotyledonous stems. Each vascular bundle is conjoint, with phloem positioned externally to the xylem, and is typically enclosed by a sheath of thick-walled sclerenchyma cells that provide mechanical reinforcement and structural integrity to the bundle. The in monocot stems forms the bulk of the stem's interior and is composed of thin-walled cells that are generally homogeneous, serving functions such as storage and metabolic support. In certain aquatic monocots, this develops extensive intercellular air spaces, forming tissue that enhances buoyancy and facilitates in submerged environments. This adaptation is particularly evident in species like those in the family , where reduces tissue density to aid flotation. Growth in monocot stems is confined to primary meristems, as the absence of secondary meristems limits radial expansion and promotes a mostly herbaceous habit. Elongation occurs through the activity of intercalary meristems located at the base of internodes or leaves, allowing for continued vertical growth even after the apex has ceased activity; this is prominently observed in grasses () and bamboos (Bambusoideae), where basal meristematic zones enable rapid height increases under favorable conditions.

Gymnosperm Stems

Gymnosperm stems exhibit distinct anatomical features adapted to their roles in production and environmental , differing from those of angiosperms through the absence of vessel elements and the presence of specialized resin-producing structures. These stems typically undergo both primary and , with the latter contributing to the characteristic woody texture observed in many species. In the primary structure of stems, a central is surrounded by primary composed primarily of tracheids, which facilitate conduction through bordered pits on their walls. Resin ducts, lined with secretory cells, are embedded within the xylem and surrounding tissues, serving as channels for flow that deters herbivores and pathogens. This arrangement is evident in young stems of , where the pith is often small and the contains additional resin canals for . Secondary growth in gymnosperm stems arises from the , producing thick layers of secondary consisting exclusively of tracheids without vessels, resulting in denser, more uniform suited for mechanical support and axial transport. The generates extensive periderm, including multiple layers of cork cells that provide robust protection against and injury. This secondary thickening allows gymnosperms to form durable trunks, with tracheids enabling efficient conduction via their pitted walls. Representative examples include such as pines (Pinus spp.), where stems bear needle-like or scale-like leaves and feature prominent resin ducts interspersed among tracheid-rich for defense against . In contrast, cycads exhibit manoxylic wood in their secondary , characterized by abundant rays and fewer tracheids, reflecting an ancient with a wide and limited woody thickening.

Pteridophyte Stems

Pteridophyte stems, found in ferns and their allies, exhibit a relatively simple vascular organization centered around the , which serves as the primary conducting tissue without the extensive secondary thickening typical of seed plants. These stems often function as rhizomes or erect structures supporting fronds, with vascular tissues arranged in patterns that reflect evolutionary advancements in complexity. The absence of a limits girth increase to primary growth and limited apical or intercalary activity. The most primitive stele type in pteridophytes is the protostele, characterized by a solid core of xylem surrounded by a cylinder of phloem, lacking any pith. This arrangement is seen in early-diverging groups such as the whisk ferns (Psilotum) and some lycophytes, providing efficient conduction in small-diameter stems. In contrast, the siphonostele represents a more advanced configuration, featuring a cylindrical ring of xylem enclosing a central pith, with phloem distributed either externally (ectophloic siphonostele) or on both inner and outer surfaces (amphiphloic siphonostele). This type occurs in marattialean ferns and some ophioglossaleans, allowing for greater structural support and accommodation of expanding pith tissue. More derived ferns, particularly in the Polypodiopsida, possess a dictyostele, where the siphonostele is fragmented into a network of interconnected vascular strands due to overlapping leaf gaps. These strands incorporate leaf traces that supply the fronds, creating a dictyostelic pattern that enhances flexibility and distribution of vascular supply in larger, more complex stems. This organization is evident in leptosporangiate ferns, facilitating the integration of reproductive and vegetative functions. In arborescent pteridophytes like tree ferns of the genus (Dicksoniaceae), the forms a pseudotrunk through persistent leaf bases and rhizome elongation, with limited thickening achieved via sclerenchyma bands in the cortical and ground tissues rather than true production. These sclerenchymatous reinforcements provide mechanical stability, supporting heights up to several meters, while the central remains dictyostelic with incorporated leaf traces. deposition in these tissues contributes to durability, highlighting adaptations for upright growth in lineages.

Stem Modifications

Storage Modifications

Plant stems have evolved various modifications to store nutrients, , and carbohydrates, enabling survival in environments with seasonal or adverse conditions. These adaptations often involve the of stems into or swollen structures that prioritize over primary functions. Such modifications are particularly common in plants, where stems serve as reservoirs during periods of or stress. One prominent type is the , a horizontal underground stem that grows parallel to the surface and stores starches and other reserves in its thickened tissues. For instance, the rhizome of ginger ( officinale) facilitates accumulation, allowing the plant to persist through unfavorable seasons and produce new shoots. Rhizomes typically exhibit nodes and internodes similar to aboveground stems but with a higher proportion of storage for efficient resource hoarding. Tubers represent another storage form, characterized by swollen, terminal portions of underground stems that accumulate large quantities of . The ( tuberosum) exemplifies this, with its developing from tips and serving as a primary site for storage, supporting regrowth after . Anatomically, tubers feature extensive parenchymal tissues that expand to maximize storage capacity while maintaining minimal vascular connections for distribution. Corms are short, vertical that are solid and thickened for , enclosed by a of dry bases. In ( spp.), the stores carbohydrates and nutrients, enabling the plant to survive and produce new shoots and from buds on its surface. Bulbs consist of short, vertical surrounded by fleshy scale leaves that together form layered organs rich in nutrients. In the onion (Allium cepa), the bulb's central stem is compact and disc-like, with the modified leaves providing the bulk of water and carbohydrate reserves. This structure allows for prolonged , with the stem base enabling the production of adventitious upon favorable conditions. In all these modifications, anatomical adaptations include a marked increase in cells, enhancing the capacity for osmotic of solutes and water. Proportionally, vascular tissues such as and are reduced to conserve space for , though they remain sufficient for minimal needs during active phases. These changes optimize allocation toward rather than . Evolutionarily, these modifications confer advantages by promoting during droughts, cold, or nutrient scarcity, while also aiding through fragmentation and resprouting. Such traits have been selected in diverse lineages, from monocots like ginger to like potatoes, enhancing in fluctuating habitats. This capacity further supports broader stem functions, such as nutrient provision during regrowth.

Reproductive and Propagative Modifications

Plant stems exhibit modifications specialized for and vegetative , enabling the production of genetically identical clones through horizontal or basal extensions that develop independent systems. These adaptations enhance dispersal and , particularly in environments where via seeds is less reliable. Key structures include stolons, offsets, and suckers, each facilitating propagule detachment and establishment via hormonal regulation and adventitious formation. Stolons, or runners, are slender, horizontal stems that extend above the surface from axillary buds at the nodes of the parent . At these nodes, adventitious emerge to anchor the developing daughter , while hormones, such as , drive root primordia initiation by promoting and elongation in the pericycle and . Once rooted, the stolon segment separates due to differential growth or mechanical breakage, forming an autonomous individual; for instance, strawberry plants (Fragaria × ananassa) utilize runners to rapidly spread and are commercially propagated by severing and planting these rooted nodes to produce uniform crops. Offsets consist of short, condensed shoots that arise laterally from the base of the parent, often with abbreviated internodes and immediate root development at the base. These structures rely on localized auxin gradients to induce adventitious roots, supplemented by cytokinins that promote shoot meristem activity for balanced growth. In houseleeks (Sempervivum spp.), offsets form dense clusters around the maternal rosette, detaching easily to colonize arid or rocky terrains through this clonal mechanism. Suckers emerge as upright shoots from adventitious buds at or below ground level, typically from a thickened rhizome-like base, and develop extensive adventitious roots influenced by signaling that overrides to allow independent establishment. Banana plants (Musa spp.) produce suckers from the corm base, which are selected and transplanted in to propagate elite varieties without . Commercially, mint (Mentha spp.) is cloned using similar runner-like extensions or suckers, harvested and rooted under controlled conditions to yield consistent profiles, leveraging node-based for efficient scaling.

Protective and Supportive Modifications

Plant stems exhibit various modifications that enhance against herbivores and environmental stresses, as well as provide for in challenging habitats. Defensive structures such as thorns primarily deter herbivory by inflicting physical damage to potential predators. Thorns are modified stems or branches originating from axillary buds, often developing into sharp, pointed structures that arise from . For instance, in ( spp.), thorns serve as rigid extensions that protect the plant from browsing animals. Associated with stems but not stem modifications themselves, spines (derived from leaves or stipules) and prickles (epidermal outgrowths without ) also provide ; for example, roses ( spp.) have prickles, while cacti like spp. have spines from areoles that reduce water loss by shading. Supportive modifications enable stems to access and stability in vertical or growth forms. Tendrils represent slender, coiling stem derivatives that actively twine around supports, facilitating upward mobility for vining plants. In grapevines (), branched stem tendrils emerge from leaf axils and exhibit , responding to touch by spiraling to anchor the plant. Cladodes, or phylloclades, are flattened, leaf-like stem expansions that provide both photosynthetic capacity and mechanical rigidity, allowing plants to withstand wind and support weight in sparse environments. species ( spp.) feature green, needle-like cladodes that replace leaves, offering structural resilience while minimizing . In arid regions, succulence emerges as a protective where stems swell to store , buffering against prolonged and enhancing survival. This modification involves enlarged cells in the that retain moisture, as seen in many cacti where succulent stems constitute the primary , enabling resilience to . Succulence thus fortifies stems against environmental extremes, complementing other defensive traits.

Functions of Stems

Mechanical Support

The mechanical support provided by plant stems ensures the elevation and positioning of leaves, flowers, and fruits for optimal light capture and , primarily through specialized s that confer flexibility, tensile strength, and rigidity. Collenchyma , composed of living cells with thickened walls, offers flexible support in young, growing stems, allowing without breakage under mechanical stress such as wind or self-weight. This is strategically located in the beneath the , providing tensile strength while permitting bending, as seen in herbaceous plants like petioles. In contrast, sclerenchyma cells, which are dead at maturity with secondary walls reinforced by , deliver rigid, long-term support through high tensile strength, resisting and in mature stems. The , particularly its lignified components, plays a crucial role in rigidity by forming a central that bears compressive loads in upright stems. impregnation in xylem cell walls creates a akin to , enabling stems to withstand gravitational forces and maintain posture. In woody stems, secondary xylem accumulates to form , which provides substantial load-bearing capacity; for instance, the modulus of elasticity in wood can exceed 10 GPa, supporting heights over 100 meters in trees like redwoods. This lignified structure integrates with to enhance overall stiffness, as referenced in discussions of development. Biomechanical adaptations further optimize support against environmental loads. Woody stems often exhibit tapering, where diameter decreases upward, distributing bending stress evenly and improving wind resistance by reducing the moment arm for lateral forces. This allometric scaling minimizes material use while maximizing stability. In large tropical trees, buttress integrate with the stem to anchor the structure against overturning moments, enhancing mechanical stability in shallow soils by increasing the root plate's . These plank-like extensions can extend several meters outward, supporting boles over 50 meters tall without deep taproots.

Fluid Conduction

The vascular system in stems, consisting of and tissues, facilitates the essential transport of water, minerals, and organic compounds throughout the body. In stems, these tissues are organized into vascular bundles, which vary in arrangement depending on the group but collectively enable efficient long-distance conduction. Xylem conducts and dissolved minerals unidirectionally from to shoots, driven by the cohesion-tension mechanism. This process, first proposed by Dixon and Joly in 1895, relies on at leaves creating negative pressure that pulls upward through conduits, with between molecules and to conduit walls maintaining continuous columns despite tensions exceeding . conduits include tracheids, found in all vascular plants, and vessels, which evolved in angiosperms and are generally more efficient for due to their wider diameter and lower resistance compared to tracheids. In angiosperms, vessels form from stacked vessel elements connected by perforation plates at their end walls, which can be simple (single large opening) or scalariform (multiple bars), reducing flow resistance and enhancing . Phloem, in contrast, transports sugars and other organic nutrients bidirectionally, primarily from photosynthetic sources to non-photosynthetic sinks. This translocation occurs via the pressure-flow hypothesis, originally formulated by Münch in 1930, where osmotic gradients generate differences that drive mass flow through conduits. sieve tubes, the primary conducting cells, are elongated and enucleate, featuring sieve plates with pores for movement, while companion cells—densely cytoplasmic and nucleated—provide metabolic support, including loading and unloading of solutes via plasmodesmata connections. This specialized structure ensures sustained translocation rates, with phloem sap velocities often reaching several centimeters per hour in stems.

Storage and Photosynthesis

Plant stems play a crucial role in resource storage, primarily through cells located in the and , which accumulate essential reserves such as , sugars, and water to support periods of growth, , or environmental stress. These thin-walled, living cells function as metabolic hubs, converting and storing carbohydrates produced during into granules that serve as reserves. For instance, in tubers—modified cells densely pack , enabling the plant to sustain and new growth from stored reserves. Similarly, the , a central region in many herbaceous stems, consists largely of storage that holds sugars and other nutrients, facilitating rapid mobilization during developmental needs. In succulent plants like cacti, stem parenchyma cells are adapted for , expanding to hold large volumes that prevent in arid environments; this hydration also supports metabolic activities by maintaining . Additionally, some parenchyma cells synthesize and store secondary metabolites, including that can act as toxins for defense against herbivores and pathogens, particularly in the transition zones of stems where such accumulation enhances protection. These storage functions are vital for survival, as they buffer against nutrient shortages or by providing on-site reserves without relying on distant production. Certain stems contribute directly to photosynthesis, especially in with reduced or absent leaves, where is embedded in the and outer layers to capture for carbon fixation. In leafless shrubs and succulents such as cacti, the surface, rich in chloroplasts within these tissues, performs the majority of photosynthetic activity, minimizing water loss through reduced surface area compared to broad leaves. Cacti exemplify this adaptation by employing the (CAM) pathway in their stems, opening stomata at night to fix CO₂ and storing it as malic for daytime use, which enhances -use in dry habitats. Some stems also utilize C4 photosynthetic pathways, concentrating CO₂ in bundle sheath-like cells to boost under high and temperature stress, as observed in non-leaf tissues of various . Hormonal regulation, particularly by auxins, orchestrates resource allocation in stems during stress, directing carbohydrates toward storage in parenchyma or modulating photosynthetic gene expression to optimize energy use. Under abiotic stresses like drought or heat, auxin signaling promotes the redistribution of reserves from source to sink tissues in stems, enhancing tolerance by prioritizing storage over growth. For example, local auxin biosynthesis and transport from storage pools in stem cells help acclimate plants to low-light or osmotic stress, fine-tuning photosynthetic capacity in chlorophyll-bearing cortex regions. This auxin-mediated control ensures balanced resource partitioning, preventing depletion of stem reserves while sustaining vital functions.

Economic and Ecological Importance

Economic Uses

Plant stems serve as a vital resource in economies, particularly through their direct utilization in food production, structural materials, and various industrial applications. Edible stems provide essential nutrition and are harvested commercially worldwide. For instance, (Asparagus officinalis) spears, which are young stems, are a low-calorie rich in , vitamins A, C, E, K, and , offering about 20 calories per half-cup serving while contributing significantly to daily vitamin requirements. , the tender stems of various species, are valued in Asian cuisines for their high protein, essential , carbohydrates, and minerals like and , with low fat content making them a healthful dietary option. () stems are primarily processed for extraction, yielding a major global source of sugar that also provides nutritional carbohydrates and antioxidants in forms like , supporting food industries and direct consumption. In materials production, woody stems from trees like (Quercus spp.) supply high-quality timber essential for , furniture, and barrels, with white oak logs commanding premium prices due to their and demand in housing and whiskey production markets. Bast fibers derived from the of (Linum usitatissimum) stems are processed into , a strong, absorbent used in and household goods, where the long fibers are separated via to create versatile yarns. Beyond food and materials, certain stems hold medicinal and value. ( spp.) rhizomes, considered , are harvested for their , which exhibit adaptogenic properties supporting immunity, cognitive function, and stress reduction in traditional and modern . Fast-growing stems of (Miscanthus x giganteus), a grass, are cultivated as feedstocks for , yielding up to 12 tons per acre annually with high energy output suitable for production and combustion.

Ecological Roles

Plant stems serve as critical microhabitats that enhance in ecosystems, particularly in forests where tree trunks and branches support epiphytic communities. Epiphytes, such as orchids, bromeliads, and lichens, colonize the rough and crevices of woody stems, utilizing them for anchorage and access to and moisture without drawing nutrients from the host. This relationship boosts overall ; for instance, in tropical rainforests, epiphyte assemblages on larger stems can comprise up to 30% of the vascular plant flora, fostering specialized food webs for pollinators, herbivores, and decomposers. In ecological succession, stems of woody plants play a pivotal role by providing structural complexity that facilitates community development. During primary succession on disturbed sites, with herbaceous or shrubby stems stabilize soil and create shaded understories, enabling the establishment of taller trees whose lignified stems form canopies in later stages. In secondary forests, the accumulation of from fallen stems further promotes by creating nurse logs that germinate seedlings and harbor fungi and invertebrates, accelerating the transition to mature climax communities. Stems contribute to nutrient cycling through and symbiotic . As stems senesce and break down, microbial activity releases stored carbon and back into the , with woody stems decomposing more slowly than leaves and thus sustaining long-term nutrient availability. Certain , like species, host in stem glands, converting atmospheric N2 into bioavailable forms that enrich surrounding soils, particularly in nitrogen-poor wetlands. In environmental interactions, stems aid and pollutant remediation. Woody stems accumulate substantial , storing carbon in lignin-rich tissues; global forests sequester approximately 7.6 billion metric tons of CO₂ annually (as of 2020), with much of the carbon stored in woody stems and tissues, mitigating through secondary thickening that locks away carbon for decades. Additionally, vascular tissues in stems facilitate by translocating and organic xenobiotics from to aboveground parts, where they accumulate or volatilize; species like willows demonstrate high stem uptake of and PCBs, reducing soil toxicity in contaminated sites.

References

  1. [1]
    Plant Development I: Tissue differentiation and function
    ... plant. Plant stem structures include: nodes, which are points of attachment for leaves and flowers; internodes, are the regions of stem between two nodes; the ...Plant Embryogenesis · Plant Body Organization · The Shoot System: Stems
  2. [2]
    Stem Anatomy - OpenEd CUNY
    The stem has three tissue systems: dermal, vascular, and ground tissue. Each is distinguished by characteristic cell types that perform specific tasks.Missing: definition | Show results with:definition
  3. [3]
    Stems - OERTX
    Their main function is to provide support to the plant, holding leaves, flowers and buds; in some cases, stems also store food for the plant. A stem may be ...Missing: definition | Show results with:definition
  4. [4]
    Vegetative plant parts | OSU Extension Service
    Their principal functions are to absorb nutrients and moisture, anchor the plant in the soil, support the stem and store food. In some plants, they can be used ...
  5. [5]
    Stems – Biology - UH Pressbooks
    Stems are part of the shoot system, providing support, connecting roots to leaves, and transporting water, minerals, and sugars. They have nodes and internodes.Missing: definition | Show results with:definition
  6. [6]
    3. Botany | NC State Extension Publications
    Feb 1, 2022 · This chapter will teach people to: Identify and describe the function of different plant parts: leaves, buds, stems, roots, flowers, fruits, ...<|control11|><|separator|>
  7. [7]
    [PDF] Plant Structures: Stems - Colorado Master Gardener
    Stems are the part of a plant that bears leaves and flowers, and they are the continuation of the vascular system pipeline that starts in the roots. Stems can ...Missing: definition | Show results with:definition
  8. [8]
    Stems - OpenEd CUNY
    Leaves are attached to the plant stem at areas called nodes. An internode is the stem region between two nodes. The petiole is the stalk connecting the leaf ...Missing: definition botany
  9. [9]
    Chapter 8: Vascular plant anatomy: primary growth - Milne Publishing
    This accounts for the presence of 'vascular bundles' in the stems of flowering plants: fundamentally they represent the traces of vascular tissue running to ...
  10. [10]
    5.2 Inside Stems – The Science of Plants
    This micrograph of a herbaceous dicot stem shows four basic parts (in order from outside to inside): epidermis, cortex, vascular bundle, and pith.Missing: botany | Show results with:botany
  11. [11]
    The Stele
    The plant stele consists of the primary vascular system of the plant axis (stem) and its associated ground tissues (eg, pith).Missing: definition | Show results with:definition
  12. [12]
    [PDF] UNIT 1: Grocery Store Botany
    Oct 8, 2012 · etymologically, the term rhizome infers a root, but botanically, a rhizome is a stem. What is going on? In this case a stem is behaving like ...
  13. [13]
    Stolon - Etymology, Origin & Meaning
    stolon(n.)​​ in botany, "a shoot, sucker," c. 1600, from Latin stolonem (nominative stolo) "a shoot, branch, sucker," cognate with Greek stēlē "upright slab," ...
  14. [14]
  15. [15]
    Chapter 1: Botany – Virginia Cooperative Extension Gardener ...
    ... plant like roots, stems, leaves, and more. Anatomy: Plant Parts and Functions. Stem Anatomy. Stems are structures that support buds and leaves and serve as ...
  16. [16]
    7.1 Meristem Morphology – The Science of Plants
    The apical meristem is the site of cell division and new cell production at the tips of the plant stems and roots.
  17. [17]
    [PDF] Vegetative plant morphology — stems, leaves, and roots
    Stems carry water and nutrients, provide framework, have xylem/phloem, pith, cambium, nodes, internodes, and leaf scars. Buds are growing points.
  18. [18]
    [PDF] Chapter 2. Vegetative morphology of plants
    The internodes are stem segments between two successive nodes. The leaves are flattened photosynthetic organs consisting of two main parts: (a) The blade (or ...Missing: botany | Show results with:botany
  19. [19]
    [PDF] THE SHOOT: ! PRIMARY STRUCTURE AND DEVELOPMENT!
    A shoot is a stem and leaves, with nodes and internodes. The apical meristem produces protoderm, procambium, and ground meristem. Stems have three basic ...
  20. [20]
    [PDF] The Shoot System I: The Stem - PLB Lab Websites
    The shoot system of a typical flowering plant consists of the stem and the attached leaves, buds, flowers, and fruits. The leaves.Missing: definition | Show results with:definition
  21. [21]
    Dicot stem
    Technically, the vascular tissues are called the stele, and stems with one ring of vascular tissues surrounding a pith are said to have a eustele. Stems as ...
  22. [22]
    Monocot stem
    Monocot stems, in which the stele consists of many vascular bundles in a complex arrangement, are said to have an atactostele.
  23. [23]
    Plant Development II: Primary and Secondary Growth
    Apical meristems contain meristematic tissue located at the tips of stems and roots, which enable a plant to extend in length. Axillary buds also contain ...
  24. [24]
  25. [25]
    Tree Rings: Recorders of Climate Change
    A full year's growth includes both a light, early wood ring and a dark, late wood ring. Counting the rings of a tree will determine its age. The pith is a ...
  26. [26]
    [PDF] Tree Growth Characteristics - Department of Plant Sciences
    In tropical climates, growth may occur year-round, and annual rings may not be visible. In some species, the rings are very pronounced (Fig. 5), because wood ...
  27. [27]
    Wood
    The cambium is a lateral (expanding in width) meristem, capable of dividing to produce additional xylem toward the inside of the stem and additional phloem to ...
  28. [28]
    [PDF] SECONDARY GROWTH IN STEMS! - The PhycoLab
    Radial system or vascular rays: ray cells formed horizontally by the ray initials, mostly ... Sapwood conducts and heartwood does not! • Older wood become ...
  29. [29]
    [PDF] Topic 05- Secondary Plant Body (Photo Atlas
    Sapwood = the outermost, generally conducting portion of the wood. Often lighter than heartwood in color. Also contains living parenchyma cells distributed as ...
  30. [30]
    [PDF] Plant Systematics Laboratory Manual
    SURFACE FEATURES. For each of the following surface types, define each character state and observe and list one example. Draw or photograph as ...
  31. [31]
    Lecture 17 - Stems primary - Daniel L. Nickrent
    Oct 14, 2022 · 1. Protostele. A solid cylinder of vascular tissue in center, found in lycophytes, ferns, stems of water plants, roots of seed plants.Missing: definition | Show results with:definition
  32. [32]
    Secondary Growth
    Cross Sections of a Woody Root: Secondary growth in the root transforms the primary structure of the organ through the formation of two cambial layers: the ...
  33. [33]
    3.3.2: Internal Anatomy of the Primary Stem - Biology LibreTexts
    Jul 28, 2025 · Primary phloem fibers cap the vascular bundles. In atactosteles (right), vascular bundles composed of xylem and phloem tissues are scattered ...
  34. [34]
    Anatomy and Primary Structure of Monocot Stem-maize Stem
    May 4, 2018 · Vascular bundles are scattered (atactostele) in the parenchymatous ground tissue. Each vascular bundle is surrounded by a sheath of sclerenchy- ...
  35. [35]
    Primary Tissues in Monocotyledonous Stem | Plants
    Vascular bundles in monocot stem usually lie scattered throughout the ground tissue and this type of distribution is termed as atactostele. In many genera like ...
  36. [36]
    Monocot and Dicot Stems - Visible Body
    In monocot stems, the vascular bundles are scattered throughout the ground tissue. Like monocot roots, monocot stems are protected by an outer layer of dermal ...Missing: atactostele | Show results with:atactostele
  37. [37]
    3 Major Groups of Plant Tissue (With Diagrams) | Botany
    In the aquatic plants, the parenchyma cells in the cortex possess well developed air spaces (intercellular spaces) and such tissue is known as aerenchyma.
  38. [38]
    Intercalary meristem - Definition and Examples - Biology Online
    Jun 16, 2022 · Intercalary meristem is a primary meristem that aids in vertical growth of plants. It's more common in monocots.
  39. [39]
    [PDF] Gymnosperms - PLB Lab Websites
    Typically, resin accumulates and flows in long resin ducts, chambers enclosed by parenchyma cells (Fig. 24.6). Resin inhibits wood-boring insects.
  40. [40]
    [PDF] Gymnosperms.pdf
    Pores that appear to be vessels are resin ducts. Close inspection will reveal that these pores are lined with parenchyma. Adjacent tracheids are connected.Missing: anatomy | Show results with:anatomy
  41. [41]
    [PDF] Evolution of development of vascular cambia and secondary growth
    Secondary growth from vascular cambia results in radial, woody growth of stems. The innovation of secondary vascular development during plant evolution ...
  42. [42]
    [PDF] Plant Anatomy - CU-PAC
    Plant anatomy is the study of plant cell and tissue structure, including the organization of the eukaryotic cell and the three tissue systems.
  43. [43]
    [PDF] IX. Seed Ferns and Cycads - Medullosales
    Wood that is rich in vascular rays is called manoxylic it is a derived wood type typical of seed ferns and cycads. These rays connect parenchyma within the ...Missing: anatomy | Show results with:anatomy
  44. [44]
    Lab VIII - Medullosans and Cycads (3)
    Their internal stem structure is characterized by a eustele with endarch protoxylem, where a small amount of manoxylic wood is produced from a bifacial ...Missing: anatomy | Show results with:anatomy
  45. [45]
    [PDF] Identifying Conifers: Arborvitae, Douglas Fir, Fir, Juniper, Pine ...
    Leaves small, scale-like, hugging the stem. Foliage in flattened plate-like display. Cones are berry-like with thick scales – Thuja (Arborvitae) – visit the Key ...
  46. [46]
    [PDF] Fabulous Ferns - UNCW
    Stele Types: Siphonosteles. • Xylem & phloem form concentric cylinders around central pith. • Ectophloic siphonostele: phloem restricted to outer surface of ...Missing: anatomy | Show results with:anatomy
  47. [47]
    [PDF] The Conservation of Artifacts Made from Plant Materials (1990)
    This stele is a protostele with a central core of xylem surrounded by phloem. Primary xylem in the stele of roots can assume different patterns. These are.
  48. [48]
    [PDF] of chicago
    In the development of the stele of Marattia alata three stages stand out sharply: (1) the protostele; (2) the amphiphloic siphono- stele, or solenostele; and ( ...
  49. [49]
    [PDF] Phylogenetic Relationships of Extant Ferns Based on Evidence from ...
    Jul 2, 2007 · UC; Bower (1923), Holttum (1964), Kaplan. (1977). 27. MATURE RHIZOME STELE TYPE: (0) protostele, (1) solenostele, (2) dictyostele, (3) eustele.
  50. [50]
    [PDF] Is Morphology Really at Odds with Molecules in Estimating Fern ...
    D 16) Mature shoot stele type: eustele (0); protostele (1); ectophloic siphonostele (2); amphiphloic siphonostele or solenostele (3); dictyostele. (4). Pryer ...
  51. [51]
  52. [52]
    What Are Strawberry Runners? (Stolons)
    Feb 11, 2022 · Strawberry plants produce runners. These stolons are horizontal stems that run above the ground and produce new clone plants at nodes spaced at varying ...
  53. [53]
    [PDF] Propogating Strawberry Plants Through Runners ... - Harvest NY
    Immediately place newly planted tips under mist propagation system. • Keep runners in a humid place. Frequent misting and humidity domes can be very helpful. • ...
  54. [54]
    How to grow sempervivum - RHS
    Learn how to grow sempervivum in your garden with the RHS expert guide on choosing, planting, feeding, pruning and propagating plants.
  55. [55]
    Plant Stem: Structure, Functions, Modifications, Facts - Microbe Notes
    May 14, 2025 · Reproduction: Certain stems reproduce vegetatively, with new plants developing through runners, suckers, and tubers. Modifications of the Stem.
  56. [56]
    Armed by Nature: Thorns, Spines, and Prickles - BYGL (osu.edu)
    Oct 4, 2023 · Rather than being made from the stuff of stems, spines are modified leaves or parts of leaves. Sometimes, spines occur in pairs, as is the case ...
  57. [57]
    [PDF] Thorns, Spines and Prickles
    Thorns and spines are modification of existing organs such as stems, leaves or stipules. Prickles or emergences are outgrowths derived from epidermal and ...
  58. [58]
    [PDF] Prickles, Thorns, and Spines, Oh My! Let's take a closer look...
    Simply put, thorns are modified branches or stems and typically end in a really sharp point. Thorns originate from the axillary bud where the leaves or branches ...
  59. [59]
    3.3.4: Stem Modifications - Biology LibreTexts
    Jul 28, 2025 · Tendrils can be derived from stems, leaves, or leaflets, and they are common in vines. Morning glory and sweet potato (Ipomoea), grapes (Vitis), ...Missing: supportive | Show results with:supportive
  60. [60]
    Acquisition and Diversification of Cladodes: Leaf-Like Organs in the ...
    This work provides evolutionary insight into the acquisition and divergence of leaf-like organs called cladodes in the genus Asparagus.
  61. [61]
    Succulent plants - ScienceDirect.com
    Sep 11, 2017 · The term 'succulence' itself is generally agreed to refer to the storage of a significant amount of withdrawable water in living cells.
  62. [62]
    Plant Cell and Tissue Types
    Collenchyma is found near the surface of cortex in stems and along the veins of leaves, where it provides structural support and protection against breakage.
  63. [63]
    6.1 Plant Cells and Tissues – The Science of Plants
    Contain a nucleus, may direct the metabolism of the sieve tube member, and are alive. Phloem fibers (sclerenchyma cells). Provide support, same as for xylem.
  64. [64]
    LON-CAPA Botany online: Supporting Tissues - Conducting Tissues
    Vascular plants have up to three types of supporting tissue: The collenchyma, a tissue of living cells,; the sclerenchyma, a tissue of nearly always dead cells, ...
  65. [65]
    Introduction to Plant Structure
    In this context, sclerenchyma provides mechanical strength to stems (fibers ... Xylem tissue functions in both water transport and mechanical support.<|control11|><|separator|>
  66. [66]
    The Role of Wood Mass Density and Mechanical Constraints in the ...
    Resistance to external wind loading can be expressed as the maximum lateral force (Fmax) at which the stress (force per area) in the stem produced by wind ...
  67. [67]
    [PDF] Size-dependent Allometry of Tree Height, Diameter and Trunk-taper
    the taper of a woody stem is influenced by the age of the stem (see Mauseth, 1988). This ontogenetic effect on taper can account for the observation that ...Missing: resistance | Show results with:resistance
  68. [68]
    An assessment of the role of buttress roots in the carbon stocks of ...
    Besides supporting and enhancing trunk mechanical stability, buttress roots ... Plate-rooted trees typically have tall trunks and canopies, with their plate roots ...
  69. [69]
    Cypress Knees: An Enduring Enigma - Arnold Arboretum
    Dec 1, 2021 · The Mechanical Support Hypothesis. Buttresses and stilt roots provide mechanical support for a number of tropical trees. It was again Lamborn ...
  70. [70]
    Water Transport in Plants: Xylem | Organismal Biology
    The cohesion-tension hypothesis is the most widely accepted model for movement of water in vascular plants. Cohesion-tension combines the process of capillary ...
  71. [71]
    LON-CAPA Botany online: Supporting Tissues - Xylem
    Wood vessels are therefore generally thought to be more efficient water conductors than tracheids. The length of the single tube (composed of numerous cells) ...
  72. [72]
    Sugar Transport in Plants: Phloem | Organismal Biology
    Phloem sap travels through perforations called sieve tube plates. Neighboring companion cells carry out metabolic functions for the sieve-tube elements and ...
  73. [73]
    Testing the Münch hypothesis of long distance phloem transport in ...
    The pressure flow hypothesis introduced by Ernst Münch in 1930 describes a mechanism of osmotically generated pressure differentials that are supposed to ...
  74. [74]
    Phloem - PropG - University of Florida
    Companion cells are parenchyma cells that function to load and unload material into the sieve tube member. Companion cells have a nucleus, while sieve tube ...
  75. [75]
    [PDF] CHAPTER OUTLINE - Plant Tissues 32 - Meristems 32
    (b) The vascular cylinder or stele in a dicot root as seen in cross section typically show the xylem in a star-shaped pattern. (c) The. Casparian strip in ...<|control11|><|separator|>
  76. [76]
    [PDF] The Organization Of The Plant Body - PLB Lab Websites
    The cortex is the region between the plant's epidermal and vascular tissues in most stems and roots. The pith is usually composed of storage parenchyma cells ...
  77. [77]
    Changes in the physiological activity of parenchyma cells in ...
    Nov 14, 2023 · Parenchyma cells biosynthesis secondary metabolites in the transition zone, and it has been suggested that the phenolics in these may be toxic ...
  78. [78]
    Pathways of Photosynthesis in Non-Leaf Tissues - PMC - NIH
    Dec 2, 2020 · The C4 pathway fixes carbon initially in a 4 carbon compound to provide a method of carbon concentration to supply the carbon required for the ...Missing: cortex cacti<|separator|>
  79. [79]
    Roles of Auxin in the Growth, Development, and Stress Tolerance of ...
    Sep 5, 2022 · This paper reviews the recent research on the role of auxin in the growth, development, and stress response of some horticultural plants.
  80. [80]
    Auxin and abiotic stress responses - PMC - PubMed Central - NIH
    Recent evidence suggests that auxin transport from local biosynthesis and storage forms plays essential roles during heat, salt, drought, and cold stresses.
  81. [81]
    A Review of the Pro-Health Activity of Asparagus officinalis L. and Its ...
    Jan 16, 2024 · Nutritional analyses have found A. officinalis to contain water (93.5%), carbohydrates (2.04%), proteins (1.91%), dietary fiber (1.31%), ...
  82. [82]
    The Nutritional Facts of Bamboo Shoots and Their Usage as ...
    Bamboo shoots have immense potential of being used as important health food as they contain high proteins, amino acids, carbohydrates, many important minerals, ...
  83. [83]
    Phytochemical profile of sugarcane and its potential health aspects
    Sugarcane crop is cultivated for the production of sugar, but the processing of sugarcane yields various valuable products such as bagasse,[8] Brown sugar ...
  84. [84]
    [PDF] the oak timber base and market - Southern Research Station
    Hardwood lumber and related sawn products are also used in the production of other goods including whiskey barrels, handles, gun stocks, solid guitar bodies and ...
  85. [85]
    Linen Most Useful: Perspectives on Structure, Chemistry, and ...
    Bast fiber, which is a major economic product of flax along with linseed and linseed oil, is described with particular reference to its application in textiles ...
  86. [86]
    Biological Activities of Ginseng and Its Application to Human Health
    Ginseng and ginsenosides seem to be beneficial for immunity, cancer, diabetes, CNS functions, and other conditions.INTRODUCTION · STRUCTURAL PROPERTIES... · ANTICARCINOGENIC...
  87. [87]
    Miscanthus Factsheet - Biofuel Decision Support System
    Giant miscanthus has strong potential to be grown as a biomass feedstock for use as a biofuel across the southeastern United States. As the bioenergy industry ...Missing: stems | Show results with:stems
  88. [88]
    Microhabitat associations of vascular epiphytes in a wet tropical ...
    Dec 5, 2014 · Microhabitat specialization of epiphyte species increased with tree size with 6% of species significantly associated with small trees and 57% ...
  89. [89]
    Epiphytic Plants: Perspective on Their Diversity, Distribution ...
    Jul 23, 2025 · Epiphytic plants are vital components of tropical and subtropical forests, contributing significantly to biodiversity, ecosystem function, ...
  90. [90]
    Distribution of Woody Plant Species Among Different Disturbance ...
    Apr 5, 2021 · Our study demonstrates the importance of forest partitioning with different disturbance regimes in maintaining local diversity in a woody plant ...
  91. [91]
    Larger fragments have more late‐successional species of woody ...
    Sep 11, 2018 · These regenerating forests are of great importance to biodiversity conservation and carbon sequestration (Barlow et al., 2007; Chazdon et al., ...
  92. [92]
    Decomposition and Carbon and Nitrogen Releases of Twig ... - MDPI
    Mar 6, 2024 · The results suggest that N inputs restrain lignin and cellulose degradation and C and N release, and increase the N/P ratio that limits P release in litter.
  93. [93]
    Nitrogen Fixation by Gunnera–Nostoc Symbiosis | Nature
    Glands occurring at the bases of leaves become invaded by the blue-green alga Nostoc puntiforme 1,2 which becomes intracellular 3 and is capable of nitrogen ...
  94. [94]
    Of Gunnera and Cyanobacteria - In Defense of Plants
    Mar 23, 2016 · From there the cyanobacteria earn their keep by producing copious amounts of usable nitrogen and in return, the Gunnera supplies carbohydrates.
  95. [95]
    Forests: Carbon sequestration, biomass energy, or both? - Science
    Incentivizing both wood-based bioenergy and forest sequestration could increase carbon sequestration and conserve natural forests simultaneously. We conclude ...
  96. [96]
    Phytoremediation | Superfund Research Center
    In this way, plants can accumulate (store) pollutants in roots, stems, and/or leaves. Some plants are very good at this, while other plants accumulate very ...
  97. [97]
    Uptake of Weathered DDT in Vascular Plants: Potential for ...
    Aug 6, 2025 · Phytoremediation is an emerging plant-based technology that may be used to cost-effectively remove or neutralize contaminants in the environment ...