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Pith

Pith is the soft, central tissue in the stems of many vascular plants, particularly dicotyledons, consisting primarily of parenchyma cells that provide structural support and nutrient storage.^[1] In some species, the pith may disintegrate over time, resulting in a hollow stem, as seen in certain canes like those of roses and grapes.^[1] This tissue is derived from the ground meristem during primary growth and is typically surrounded by vascular tissues in a cylindrical arrangement.^[2] In plant anatomy, pith occupies the innermost region of the stem, distinct from the surrounding cortex and vascular bundles, and its cross-sectional appearance—such as solid, chambered, or star-shaped—serves as a key identifier in botanical studies.^[1] Composed of thin-walled parenchyma cells capable of storing starch, sugars, and water, pith contributes to the plant's metabolic reserves and can exhibit totipotency, allowing regeneration of other cell types if damaged.^[2] While most prominent in dicot stems, pith also occurs in the central region of some roots, where it functions similarly as a storage site.^[2] In woody plants, the pith persists as a small core within the wood, highlighting its role in early developmental stages before secondary growth dominates.^[3]

Botany

Definition and structure

Pith is the central, soft, spongy parenchyma tissue located in the stems and roots of many vascular plants, particularly eudicots and pteridophytes.^[2]^[4] It forms the innermost portion of the primary plant body, providing a foundational ground tissue that contrasts with the more specialized vascular and dermal layers.^[5] Microscopically, pith is composed primarily of parenchyma cells with thin, primary cell walls, large central vacuoles that occupy much of the cell volume, and extensive intercellular spaces that impart its characteristic sponginess.^[6]^[7] These cells are typically alive at maturity, with protoplast pressed against the walls by the vacuoles, enabling flexibility and potential for regeneration.^[8] In certain species, the pith may incorporate sclereids or fibers, which are thickened cells that enhance mechanical support without compromising the overall softness.^[8] The pith differs markedly from adjacent stem and root tissues in both position and role within the tissue system. It occupies the core, surrounded by the vascular cylinder (stele), whereas the xylem facilitates water and mineral transport through lignified conduits, the phloem distributes sugars via sieve elements, and the cortex acts as an external ground tissue layer for storage and protection just beneath the epidermis.^[5]^[2] Developmentally, pith originates from the ground meristem, one of the three primary meristems arising from the apical meristem during primary growth, and is situated as the innermost region of the stele.^[2] This derivation positions it centrally within the differentiating embryo and seedling, where it expands through cell division and elongation to fill the core of the axis.^[9]

Occurrence in plants

Pith is prominently featured in the stems of eudicots, where it occupies the central region as a mass of parenchyma cells, as seen in herbaceous species like elderberry (Sambucus spp.), whose young stems contain soft, white pith that can be easily hollowed.^[10] In citrus plants (Citrus spp.), pith occurs in stems, serving as a structural component in young growth.^[11] It is also common in herbaceous dicots, forming a distinct core surrounded by vascular bundles arranged in a ring.^[11] In gymnosperms, pith occurs but is often limited in extent, particularly in conifers such as pine (Pinus spp.), where it appears as a small central region of parenchyma near the primary xylem in young stems.^[12] The pith persists throughout the tree's life in many conifers, though it may become compressed or reabsorbed in some species like those in Araucariaceae and Podocarpaceae.^[13] Pith is reduced or absent in monocots, where stems typically lack a central pith region due to the scattered arrangement of vascular bundles throughout the ground tissue.^[14] In woody plants, pith often diminishes with secondary growth as the vascular cambium expands, producing layers of secondary xylem and phloem that push the pith inward to form a small, residual core.^[3] For instance, in trees like apple (Malus spp.), the pith remains a narrow central cylinder within the wood after extensive secondary thickening.^[15] Pith is absent in non-vascular plants such as mosses and algae, which lack true stems and vascular tissues altogether.^[16] In ferns, pith is present as a central medulla within siphonosteles, where vascular tissue forms a cylinder around the pith-filled core.^[17] Similarly, horsetails (Equisetum spp.) feature a central pith or medulla cavity surrounded by vascular bundles in their jointed stems.^[18] Evolutionarily, pith represents a primitive feature in vascular plants, originating from early protosteles and retained primarily in younger stems for space-filling and structural support before secondary growth predominates in more derived lineages.^[19]

Physical properties

Pith exhibits a soft, spongy texture owing to its composition of thin-walled parenchyma cells with abundant air-filled intercellular spaces, resulting in a low relative density typically around 0.01 in representative examples like potato parenchyma.^[20] This structure imparts a foam-like quality, with the tissue appearing white or pale in color due to the absence of significant pigmentation in the cell walls.^[20] Moisture content in pith is high, generally ranging from 70% to 90% of fresh weight, which contributes to its pliability and hydration-dependent behavior in plant stems.^[21] Mechanically, pith is highly compressible and absorbent, with low tensile strength that allows it to deform under minimal stress without fracturing; for instance, Young's modulus values fall between 0.3 and 14 MPa in parenchyma tissues such as those in apple and potato.^[20] Its compressive strength is similarly modest, around 0.25 to 1.3 MPa, enabling expansion upon hydration—up to several times the original volume in absorbent species like elder (Sambucus nigra), where dried pith can swell significantly when rewet.^[20] These properties stem from the thin cell walls, which lack lignification in young tissues, providing elasticity rather than rigidity.^[20] Chemically, pith is dominated by water, alongside starch for storage, mucilage polysaccharides that enhance hydration capacity, and a matrix of cellulose, hemicellulose, and pectin in the primary cell walls, comprising about 33% pectin in some cases.^[20] The tissue maintains a pH that is neutral to slightly acidic (around 5-6), reflecting the metabolic environment of living parenchyma, and is generally non-toxic, though it may contain tannins in certain species that impart mild astringency.^[21] Pith properties vary with plant age: in younger stems, the tissue remains elastic and hydrated due to active parenchyma cells, but as plants mature, progressive lignification of cell walls increases density and stiffness, rendering older pith more brittle and less absorbent.^[20]^[22] This transition enhances mechanical support in woody species but reduces the spongy character observed in herbaceous plants.^[23]

Biological functions

Role in growth and development

The pith originates during early plant embryogenesis as part of the ground meristem, a primary meristem that differentiates into the ground tissue system of the shoot.^[24] This meristematic layer forms the central core of the stem, consisting primarily of parenchyma cells that establish the foundational structure before vascular tissues fully develop.^[25] In herbaceous and young perennial plants, the pith persists as a distinct tissue, whereas in woody species undergoing secondary growth, it is often compressed or obliterated by expanding secondary xylem.^[26] During primary growth, the pith serves as a lightweight filler in young stems, composed of thin-walled parenchyma cells that prevent structural collapse until the vascular tissues, such as xylem and phloem, mature and provide rigidity.^[27] Its low-density composition allows for efficient resource allocation, enabling stems to thicken with minimal investment in dense supportive tissues like wood.^[28] Additionally, the pith contributes to turgor pressure through water retention in its vacuolated cells, which maintains stem uprightness and elasticity during elongation. In herbaceous plants, the pith plays a key role in accommodating rapid developmental changes. For instance, in sunflowers (Helianthus annuus), the extensive pith forms the bulk of the stem's interior, offering lightweight support that facilitates quick vertical elongation while the peripheral vascular bundles handle primary mechanical strength.^[29] Similarly, in flexible vines such as grape (Vitis vinifera), the pith's parenchyma cells with thin, pliable walls enhance stem suppleness, allowing bending without breakage during growth and environmental adaptation.^[30] Post-formation, pith cells exhibit limited mitotic division but undergo significant radial expansion to accommodate stem widening during primary growth.^[31] This expansion is particularly evident in the transition to secondary growth, where pith ray cells may integrate with the emerging vascular cambium, facilitating the shift from primary elongation to girth increase in perennials.^[32] In woody plants, this interaction often results in the pith being progressively compressed as secondary tissues accumulate.^[26]

Storage and defense mechanisms

The pith functions as a key storage tissue in many plants, accumulating starch, sugars such as fructans, ions, and water to serve as reserves for periods of dormancy or stress. In herbaceous stems and roots, its parenchyma cells act as a carbohydrate sink, particularly during dormancy, enabling sustained growth upon reactivation; for instance, in Jerusalem artichoke (Helianthus tuberosus) tubers, which are modified underground stems, the parenchyma stores high levels of inulin—a fructan polymer—comprising up to 75% of dry weight as an energy reserve for spring regrowth.^[33]^[34] Water storage is prominent in pith parenchyma of certain stems, such as in sorghum (Sorghum bicolor), where living cells maintain high juice content (up to 8 times more than in dry-stem varieties) to support metabolic processes like sugar production.^[35] In addition to nutrient and water reserves, the pith contributes to plant defense through the production of secondary metabolites that deter herbivores and pathogens. Upon herbivore attack, such as by stem-boring weevils (Trichobaris mucorea) in Nicotiana attenuata, pith tissue locally accumulates phenolic compounds like chlorogenic acid (CGA), increasing levels up to 1000-fold via jasmonic acid signaling to inhibit larval development and survival without systemic effects on other herbivores.^[36] In citrus species, the albedo (pith layer) concentrates limonoids—bitter triterpenoids such as limonin—that act as feeding deterrents and growth disruptors against insect pests, enhancing fruit protection.^[37] These localized responses minimize resource allocation to non-attacked tissues, promoting efficient defense. The pith also plays a role in wound response and tissue repair, facilitating regeneration after injury. In tobacco (Nicotiana tabacum), wounded pith cells redifferentiate by resuming division to form a wound meristem, producing callus tissue that differentiates into vascular elements like tracheary cells and cambium, thereby sealing damage and restoring continuity; this process is auxin-dependent, with higher concentrations expanding the regenerative zone.^[38] Ecologically, pith storage enhances plant survival in nutrient-poor or stressful environments by enabling internal recycling of reserves during periods of limited external uptake. In high-altitude species like Espeletia (Andean rosette plants), large pith reserves sustain transpiration and growth when frozen soils restrict root absorption, supporting adaptation to oligotrophic conditions.^[39] This buffering capacity contributes to community resilience in resource-scarce habitats by allowing nutrient remobilization to reproductive or vegetative sinks under drought or nutrient stress.^[40]

Human uses

Culinary applications

Pith from various edible plants serves as a versatile ingredient in culinary preparations, particularly when processed to mitigate its natural bitterness and potential toxicity. In citrus fruits such as oranges and grapefruits, the white pith underlying the peel is commonly incorporated into marmalades, where it provides natural pectin for gelling the preserve.^[41] This practice leverages the pith's high pectin content, which is concentrated in the albedo layer, enabling the mixture to set without additional thickeners.^[42] Similarly, the tender inner pith of young bamboo shoots is a staple in Asian cuisines, featured in stir-fries, soups, and curries after initial boiling to neutralize cyanogenic glycosides like taxiphyllin.^[43] Young shoots from elderberry plants can also be peeled and cooked like asparagus in some traditional foraging recipes, though stems must be thoroughly boiled to eliminate toxins.^[44] Preparation methods for pith emphasize reducing its inherent bitterness and ensuring safety. Boiling is a standard technique for bamboo shoots and elderberry stems, which not only softens the texture but also degrades cyanogenic compounds, rendering them safe for consumption—typically requiring 20-30 minutes at high heat.^[43] For citrus pith, candying or prolonged simmering with sugar balances the astringency, as seen in marmalade recipes where the pith is sliced thinly and cooked for hours.^[45] The mucilaginous quality of pith, derived from soluble fibers like pectin, also functions as a natural thickener in dishes such as soups, enhancing mouthfeel without synthetic additives.^[42] Nutritionally, plant pith offers benefits centered on its fiber content, with citrus varieties providing soluble pectin that supports digestive health by promoting gut regularity and potentially lowering cholesterol absorption.^[46] Bamboo shoot pith is low in calories (approximately 27 kcal per 100 g) and rich in dietary fiber (around 2 g per 100 g), contributing to satiety and weight management, while also supplying modest amounts of vitamin C and B vitamins.^[47] However, unprocessed pith from certain sources, such as bamboo or cassava-related plants, may harbor cyanogenic glycosides that necessitate thorough cooking to prevent cyanide release, underscoring the importance of proper preparation.^[43] Culturally, pith has featured in diverse traditions, including Chinese cuisine where dried bamboo pith (often from fungal growths on bamboo) is rehydrated and added to nourishing soups for its subtle texture and purported health benefits.^[48] In Europe, the use of citrus pith in jams dates to the 18th century, when Scottish producers like the Keillers commercialized orange marmalade, transforming the bitter component into a breakfast essential that balanced tartness with sweetness.^[49] These applications highlight pith's role in both everyday meals and preserved foods across continents.

In headwear and materials

Pith from the stems of Aeschynomene aspera, commonly known as sola or shola, has been a primary material for crafting lightweight protective headgear, particularly the sola topi or pith helmet, designed for sun protection in tropical climates. The plant's spongy, milky-white pith is harvested from mature stems, which are cut and the outer layers removed to extract the soft core. This pith is then dried, pressed into thin sheets, and laminated over dome-shaped molds to form the helmet's rigid structure, resulting in a product weighing typically under 200 grams that provides effective heat insulation due to its low thermal conductivity.^[50] The formed helmet is coated with shellac for durability and water resistance, followed by covering in fabric such as khaki or white cotton, with ventilation holes and a wide brim added for enhanced airflow and shade.^[51] These pith helmets gained prominence in British India during the 19th century, first appearing in the 1840s during the Anglo-Sikh Wars and becoming standard issue after the Indian Mutiny of 1857 for colonial officials, military personnel, and civilians to shield against intense solar radiation.^[52] The sola topi's buoyancy and insulating qualities also made it suitable for use near water or in humid environments, contributing to its adoption across British colonial territories in Africa and Asia.^[53] In modern times, variants persist as safari hats or decorative items, retaining the original lightweight design for outdoor activities in hot regions.^[54] Beyond headwear, sola pith's compressible and buoyant nature has found applications in other lightweight materials, such as insulation panels and flotation devices, leveraging its low density for thermal and buoyant properties.^[55] Elder (Sambucus nigra) pith, being soft and easily removable from stems, has been historically used in crafts for creating lightweight models, beads, and hollow tubes like whistles or pipes, valued for its workability in small-scale decorative and functional items.^[56]

Cleaning and maintenance

In horology, elder pith (Sambucus nigra) is cut into small pegs or sticks to gently remove oil, dust, and residues from watch mechanisms, leveraging its soft texture to avoid scratching delicate parts while polishing pivots to a fine finish.^[57]^[58] This non-abrasive quality stems from the material's low density and spongy parenchyma cells, making it ideal for precision tasks.^[59] The use of elder pith in cleaning became a standard practice in 18th- and 19th-century clockmaking, as detailed in period horological manuals, and continues today among professional restorers for maintaining intricate gears and escapements without risking damage.^[57]^[60] Modern suppliers provide pre-cut bundles of natural elder pith sticks, typically 80–100 mm long, for this purpose.^[61] Elder pith also finds application in cleaning scientific instruments, such as microscopes and optical components, where it is shaped into swabs to absorb and extract contaminants without leaving fibers or abrasives.^[62] Its inherent absorbency effectively draws out lubricants and impurities, ensuring clarity in sensitive optics.^[59] Preparation involves harvesting young elder stems, extracting the central pith, and forming it into usable shapes; dried pith is most common for durability and availability, though it maintains pliability when stored properly.^[59]^[57]

As an illuminant

Pith from various plants has been employed historically as a wick material in rudimentary candles known as rushlights, particularly in medieval Europe where the pith of rushes such as Juncus effusus was stripped of its outer skin, peeled into thin strips, and soaked in tallow or animal fat to create inexpensive lighting for households.^[63]^[64]^[65] These rushlights provided a basic, clear flame but required frequent adjustment, burning for approximately 30 minutes per 15-inch length when held at an angle in simple holders.^[66] In Asia, similar techniques persisted; Japanese candles traditionally incorporated rush pith wicks alongside paper for oil lamps, while in India, fibers extracted from banana stem pith (Musa spp.) were rolled into wicks for oil lamps used in rituals and daily illumination, valued for their even burning due to the material's porosity.^[67]^[68] Beyond wicks, dry pith from plants like elder (Sambucus nigra) and mullein (Verbascum thapsus) served as tinder in fire-starting applications across Europe and North America, prized for its spongy texture that readily caught sparks from flint and steel with minimal smoke and rapid ignition.^[69]^[70]^[71] This use dated back to Anglo-Saxon times for elder and extended into pioneer practices, where the pith's low density allowed it to propagate embers effectively for kindling larger fires.^[72] In modern contexts, pith-based illuminants persist in survival scenarios and artisanal recreations, where mullein or elder pith tinder facilitates primitive fire lighting, and banana pith wicks appear in traditional Indian lamps for cultural ceremonies, leveraging the material's ability to absorb and distribute oil evenly for sustained, low-flame burning.^[71]^[68] However, these applications were largely supplanted by the mid-19th century with the advent of paraffin wax candles, which offered longer burn times and cleaner light, rendering pith wicks obsolete for widespread use.^[73]^[74]

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To clean the face and ascertain if it is well polished, a small piece of bread or elder pith is taken in which the pinion is stuck, making sure that the ...
[62]
https://darwin-online.org.uk/converted/pdf/1855_Schacht_microscope_DlibD_A5802.pdf
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[PDF] The Microscope - Darwin Online
atmospheric deposits, it may be cleaned with dry elder pith, care being taken to cut off with a clean razor the surface of the pith which has once been used ...
[64]
Out of the Dark - John Moore Museum
Jul 27, 2021 · Rushes were gathered and dried out and then stripped of their skin, this revealed the inner pith which served as a basic wick.Missing: plant | Show results with:plant
[65]
18th and 19th Century Lighting – North Haven Historical Society and ...
Rush lights were the simplest form of candle. The pith, or core, of rushes or reeds were dried, then saturated with waste kitchen fat and held in rush light ...
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Rush Lights: Illuminating the Dark Ages - - Medieval Historia -
Jun 24, 2025 · They consisted of the pithy stem of a rush plant, typically the common soft rush (Juncus effusus), which was soaked in fat or oil and then lit.Origins Of Rush Lights · Making Of A Rush Light · Reasons For Using Rush...
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Rushlight: How the Country Poor Lit Their Homes - The Victorian Web
Feb 1, 2009 · About an inch and a half at a time was pulled up above the jaw of the holder. A rush-light fifteen inches long would burn about half-an-hour.
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The Japanese Craftsman | Technology Magazine Metamorphosis
The wicks of Japanese candles are made of traditional Japanese paper and rushes. The rush plant grows in marshy areas and shallow water, and is used to make ...
[69]
Novel banana core stem fiber from agricultural biomass for ...
Artifacts such as candle wick, oil lamp wick and ear buds have been made with these fibers. The carbon deposit by the candle wick made with the banana core stem ...Missing: historical | Show results with:historical
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Elder guide: how to identify, when and how to forage for elderflowers ...
May 22, 2022 · Later still, the Anglo-Saxons gave their name for fire, aeld, to elder, perhaps because the pith could be dried as tinder. Elder, however ...
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Elderberry – A “Miracle Plant”? | The Granada Native Garden ...
Aug 23, 2013 · The soft pith from the inside of the branches could be used as tinder for starting fires. The genus name, Sambucus, happens to be taken from the ...
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Natural Plant Tinders - Wilderness Solutions
The pith of the mullein weed stalk may require two or three attempts before lighting. Once the ember has formed however it propagates well and can be used to ...
[73]
Elderberry a versatile plant throughout history - Naturally North Idaho
Aug 6, 2015 · The pithy stems were hollowed out for flutes and whistles with the pith being used as tinder. Since the plant grows in clumps, 10 to 30 stems ...
[74]
Candles In The Victorian Age: Looking Back To Look Forward
May 10, 2017 · Patented in 1850, paraffin wax was created after chemists in Battersea, UK, discovered how to efficiently separate this natural byproduct of ...
[75]
History of Candles
Historians note that the first “standard candles” were made from spermaceti wax. 19th Century Advances. Most of the major developments impacting contemporary ...