Vegetative Reproduction

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Vegetative reproduction is a type of asexual reproduction in plants where a new individual is formed from a multicellular part of the vegetative body of the mother plant (shoot, leaf, root) (Serebryakova et al., 2006; Yakovlev et al., 2008). In contrast to true asexual reproduction, which is carried out by specialized cells – spores – vegetative reproduction occurs via somatic (body) cells that have already undergone differentiation but retain the ability to restore a whole organism.

The essence of the process is the detachment of a viable portion of vegetative organs (shoot, root, leaf) from the mother plant and subsequent regeneration from it of a complete daughter individual, genetically identical to the mother. The totality of all plants originating from one initial individual by vegetative reproduction is called a clone (Serebryakova et al., 2006).

Biological basis: totipotency and regeneration. The ability for vegetative reproduction is based on two fundamental properties of plant cells:

Totipotency – the ability of any living somatic cell (even a highly specialized one) to realize all the genetic information contained in the nucleus and give rise to a whole plant (Bidabadi & Jain, 2020). Thanks to totipotency, a full-fledged organism can develop from a fragment of a stem, leaf, or root.
Regeneration – the ability of a plant to restore lost parts of organs or a whole plant from its fragment. In vegetative reproduction, regeneration manifests itself in the formation of adventitious roots on stem cuttings, adventitious shoots (root suckers), or the formation of a whole plant on a leaf (Garcès et al., 2007; Bell, 1991).

The regeneration process includes metabolic reprogramming of cells in the wound zone, their dedifferentiation (return to a meristematic state), callus formation or direct differentiation of new tissues and organs (Bidabadi & Jain, 2020).

Place in the plant reproduction system. In the plant reproduction system, vegetative reproduction occupies an intermediate position between true asexual (spore) and sexual reproduction.

In sexual reproduction, gametes fuse to form a zygote carrying a recombined genotype from both parents. This provides genetic variability but requires energy expenditure for flower formation and pollinator attraction (Evert & Eichhorn, 2013).

In asexual (spore) reproduction (characteristic of ferns, mosses, horsetails), daughter individuals arise from haploid spores formed in sporangia by meiosis. This form of reproduction also leads to dispersal but often involves alternation of generations (sporophyte and gametophyte) (Serebryakova et al., 2006).

Vegetative reproduction is not associated with spore or gamete formation. It yields exact genetic copies (clones) of the mother plant, which is of immense importance for preserving valuable cultivar traits in agriculture (Carmel et al., 2024). However, the lack of genetic recombination makes clones vulnerable to diseases and environmental changes (Evert & Eichhorn, 2013).

Natural and artificial vegetative reproduction. Two forms of vegetative reproduction are distinguished:

Natural – occurs in nature without human intervention. Examples: reproduction by stolons (strawberry), rhizomes (couch grass, iris), tubers (potato), bulbs (tulip, lily), brood buds (bryophyllum, viviparous knotweed), root suckers (aspen, cherry, raspberry) (Serebryakova et al., 2006; Yakovlev et al., 2008). Natural vegetative reproduction is often an important mechanism for territory conquest and survival under conditions where seed regeneration is difficult (Tan et al., 2025).
Artificial – carried out by humans to propagate economically valuable plants. Main methods: cutting, layering, division of the bush, grafting, and micropropagation (tissue culture) (Hartmann et al., 2011; Carmel et al., 2024). Artificial vegetative reproduction allows obtaining a large amount of genetically uniform planting material in a short time.

Thus, vegetative reproduction is an effective strategy for preserving and reproducing a genotype, based on the remarkable ability of plants to regenerate from somatic cells. Understanding its essence is necessary for moving on to the study of specific mechanisms, classification of types, and practical techniques for managing this process in agronomy.

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1. Biological and evolutionary significance

Vegetative reproduction is not just a method of reproduction but an entire evolutionary strategy that has its own biological “pros” and “cons” compared to seed (sexual) reproduction. Understanding these advantages and limitations is necessary for the proper use of vegetative reproduction in agronomy and breeding.

Biological significance: advantages and disadvantages

Advantages of vegetative reproduction

Exact preservation of the genotype (cloning). In vegetative reproduction, daughter individuals are exact genetic copies of the mother plant. This is especially valuable if the mother plant possesses a unique combination of economically valuable traits (high yield, disease resistance, special taste qualities). In seed reproduction, these traits segregate in the progeny due to genetic recombination (Evert & Eichhorn, 2013; Carmel et al., 2024). This is why most varieties of fruit, berry, and ornamental crops are propagated vegetatively.
Preservation of heterosis effect. Many high-yielding varieties are interspecific or intervarietal hybrids. In the first generation, such hybrids exhibit heterosis (“hybrid vigor”) – superiority over parents in several traits. However, in sexual reproduction, heterosis is lost in subsequent generations. Vegetative reproduction allows fixing the heterosis effect indefinitely (Sakthivel et al., 2025). A classic example is the cultivated banana, which is propagated only vegetatively (by rhizomes and basal suckers), as its seeds are sterile.
Rapid biomass accumulation and territory conquest. Vegetative reproduction often occurs much faster than seed reproduction. The resulting daughter individuals (ramets) already have formed vegetative organs and can immediately start photosynthesis and nutrition, while a seedling needs time for germination and establishment (Tan et al., 2025). This gives vegetatively reproducing plants a competitive advantage when colonizing new habitats, which is especially evident in invasive species (e.g., Canadian waterweed, alligator weed).
Ability to reproduce in the absence of pollinators or mates. For many plants, vegetative reproduction is a “backup” option when seed reproduction is impossible for some reason. This may be due to the absence of insect pollinators, unfavorable weather conditions during flowering, or the fact that the population contains individuals of only one sex (as in dioecious Canadian waterweed, introduced to Europe only as female specimens). In such cases, vegetative reproduction becomes the only way to maintain the population (Serebryakova et al., 2006; Yakovlev et al., 2008).
Rejuvenation and dispersal. Many plants form specialized organs of vegetative reproduction (brood buds, bulbils, tubercles) that easily detach and are dispersed by wind, water, or animals. Such diaspores often have a much younger (rejuvenated) physiological age than the mother plant, ensuring high viability and growth energy (Garcès et al., 2007). Example: bryophyllum (kalanchoe) forms entire plantlets with roots on the leaf margins, which quickly root after falling off.

Disadvantages and limitations

Genetic uniformity (lack of variability). This is the main biological drawback. Since all individuals of a clone are identical, they are equally sensitive to unfavorable environmental factors, diseases, and pests. An outbreak of disease or the emergence of a new pest can destroy the entire clone. A classic historical example is the “Irish potato famine” of 1845–1852, when late blight destroyed almost the entire potato crop, which was represented by only a few poorly resistant cultivar clones (Evert & Eichhorn, 2013). Genetic diversity created by sexual reproduction serves as an “insurance” against such catastrophes.
Pathogen accumulation. During prolonged vegetative reproduction, viruses, bacteria, and fungi accumulate in plant tissues, being transmitted from the mother plant to all offspring. This leads to gradual cultivar degeneration – reduced yield, product quality deterioration, and weakened growth. This is why regular sanitation of planting material (micropropagation, thermotherapy) is necessary in fruit growing and potato production (Sakthivel et al., 2025).
Lack of long-distance dispersal. Although some diaspores of vegetative origin can be carried over considerable distances, vegetative reproduction generally provides mainly local (short-range) territory conquest. Seeds, especially those equipped with adaptations for anemochory or zoochory, can travel tens and hundreds of kilometers, allowing a species to rapidly colonize new areas (Yakovlev et al., 2008).

Evolutionary significance

From an evolutionary point of view, vegetative reproduction is not a “dead end” but an important adaptive mechanism that has arisen multiple times in different plant groups.

Role in life cycles. Many plants exhibit a combination of vegetative and seed reproduction. For example, strawberry reproduces both by stolons (cloning) and by seeds (recombination). Such a “flexible” strategy allows the species to simultaneously preserve successful genotypes and create new gene combinations for adaptation to changing conditions (Evert & Eichhorn, 2013).

Evolution of facultative asexuality. Studies on duckweed Spirodela polyrhiza have shown that in populations with predominance of vegetative reproduction, genetic diversity and DNA methylation levels (in particular, CHH methylation) are reduced (Wang et al., 2024). Interestingly, natural selection in such populations often acts on genes controlling the transition to flowering and seed reproduction, which may lead to the “fixation” of the vegetative strategy.

Emergence of specialized organs. The evolution of vegetative reproduction has led to the appearance of amazing morphological structures: tubers, bulbs, rhizomes, brood buds, and even entire “baby plants” on leaves (as in kalanchoe). Genetic analysis has shown that the formation of these structures is often associated with the “rewiring” of genes that in ancestors were responsible for embryogenesis (seed embryo development) (Garcès et al., 2007). For instance, in kalanchoe with constitutive (constant) formation of brood buds on leaves, the LEC1 gene, critically important for normal embryo development and seed desiccation tolerance, turned out to be defective. This led to seeds becoming non-viable in such species, and the reproductive function shifted entirely to vegetative structures (Garcès et al., 2007).

Thus, vegetative reproduction is not a primitive but a highly adaptive and evolutionarily plastic mode of reproduction. It allows a plant to:

fix successful genotypes under stable conditions;
rapidly conquer space and form dense clumps;
survive where seed reproduction is difficult or impossible.

However, the price for these advantages is genetic uniformity and susceptibility to pathogens, which in the long term limits the evolutionary potential of purely vegetative lineages. Therefore, species that combine both modes of reproduction – sexual and vegetative – dominate in nature.

2. Anatomical and physiological basis of vegetative reproduction

To understand why a whole plant can grow from a piece of stem or leaf, it is necessary to understand the anatomical structures and physiological processes underlying regeneration. Key roles here are played by meristems, polarity, and the ability of cells to reprogram.

Role of meristems

Meristems are tissues composed of undifferentiated (or weakly differentiated) cells that retain the ability to divide actively. They are responsible for plant growth and regeneration.

In the context of vegetative reproduction, three types of meristems are important:

Apical (terminal) meristems. Located at the tips of shoots and roots. Their division ensures organ elongation. In vegetative reproduction by shoot fragments (cuttings), the apical meristem is preserved and continues development, forming a new above-ground part (Serebryakova et al., 2006).
Axillary (lateral) meristems. Located in leaf axils and give rise to lateral shoots. In cuttings or layering, these meristems are often activated and become the basis for forming a new crown (Bell, 1991).
Adventitious meristems. Form endogenously (from deep tissues) where they are not normally found. They can arise from cambium, pericycle, phloem, parenchyma cells. It is adventitious meristems that give rise to adventitious roots on stem cuttings and adventitious shoots on root suckers (Bell, 1991; Evert & Eichhorn, 2013).

Vegetative reproduction is often associated with the presence of specialized organs containing meristem-rich tissues: tubers (eyes are axillary buds), bulbs (basal plate – a shortened stem with numerous meristems), rhizomes (nodes with axillary buds). In these organs, meristems are dormant and start growing when favorable conditions arrive.

Cell dedifferentiation and totipotency

Diagram: from zygote to plant and from differentiated cells to callus and embryoids — CC BY 4.0
Totipotency and pluripotency in plant regeneration
The figure shows how somatic cells can dedifferentiate and regain the ability to form a whole organism.

When a plant part (cutting, leaf) is detached, cells in the cut zone find themselves under stress: tissue integrity is disrupted, vessels are damaged, hormonal balance changes. In response to wounding, many differentiated cells (parenchyma, phloem, and even xylem cells) lose their specialized features and return to a state close to meristematic. This process is called dedifferentiation (Bidabadi & Jain, 2020).

Dedifferentiated cells acquire totipotency – the ability to realize the entire genetic potential of the nucleus and give rise to any type of cells and tissues, and ultimately to a whole organism. It is totipotency that underlies all methods of plant regeneration from somatic cells.

Dedifferentiated cells actively divide and form callus – a loose mass of unorganized parenchyma cells arising on the wound surface. Callus has no tissue structure, but its cells retain high meristematic activity. Under certain conditions (with an appropriate hormonal balance), new meristematic foci are established in the callus, which then differentiate into adventitious roots, stems, or even embryo-like structures – somatic embryoids (Bidabadi & Jain, 2020).

It is important to note that callus can form not only at the cut site but also over the entire surface of a damaged organ. This property is widely used in biotechnology for micropropagation and production of somatic hybrids.

Polarity

Polarity is a fundamental property of plants, manifested in the clear spatial organization of axes (shoot – root, top – bottom). Even at the level of an isolated cutting or tissue fragment, polarity is preserved.

Classic experiment: a willow cutting placed in a humid environment always forms roots at its basal (lower, closer to the original plant’s roots) end, and shoots at its apical (upper, farther from the roots) end (Evert & Eichhorn, 2013). If the cutting is planted “upside down” (apical end in the substrate), roots still appear at the end that was basal, even if that end is in the air. Polarity does not depend on external factors (light, gravity) and is determined by the internal structure of vascular bundles and the distribution of phytohormones, primarily auxins.

The mechanism of polarity in cutting rooting is explained by polar auxin transport: auxin is synthesized in young leaves and the apical meristem and moves strictly basipetally (from tip to base). In the lower part of the cutting, auxin accumulates and stimulates adventitious root formation (Pincelli-Souza et al., 2024). This is the basis for treating cuttings with synthetic auxins (IBA, NAA) – they enhance the natural gradient and accelerate rhizogenesis.

Polarity is preserved even in isolated tissue pieces and cell cultures, which must be taken into account in micropropagation and somatic embryogenesis.

Apical dominance

Apical dominance is the phenomenon where the apical bud (apical meristem) suppresses the growth of lateral (axillary) buds. It is caused by the apical meristem producing auxin, which inhibits the development of lower buds.

In vegetative propagation by cuttings, the apical bud is often removed (or the cutting is taken without it). This releases apical dominance, and lateral buds begin to develop actively, forming a branched above-ground part (Carmel et al., 2024). In horticultural practice, to stimulate bushing and obtain compact seedlings, the apical bud of a cutting or grafted plant is sometimes pinched off (removed).

On the other hand, in propagation by root suckers or rhizomes, apical dominance is weakly expressed because buds are formed on roots or underground shoots where auxin concentration is lower and cytokinin concentration is higher.

Thus, the anatomical and physiological basis of vegetative reproduction includes:

the presence or ability to form meristems (apical, axillary, adventitious);
dedifferentiation of cells and callus formation;
totipotency of somatic cells;
polarity, which determines the directionality of organogenesis;
apical dominance, regulating the balance between main shoot and lateral shoot development.

Understanding these fundamental processes is necessary for developing effective methods of artificial vegetative reproduction – from traditional cutting to modern biotechnological techniques.

3. Classification of types of vegetative reproduction

The diversity of forms of vegetative reproduction is usually divided into two large groups: natural (occurring in nature without human intervention) and artificial (carried out by humans for economic purposes). Within each group, methods are distinguished based on which vegetative organ (shoot, root, leaf) or its modification serves to produce offspring.

3.1. Natural vegetative reproduction

In natural conditions, vegetative reproduction ensures species survival, rapid colonization of territory, and preservation of successful genotypes. It is carried out using specialized or non-specialized parts of the plant.

Reproduction by modified shoots

By rhizomes. A rhizome is a perennial underground shoot with nodes, internodes, scale-like leaves, and axillary buds. Adventitious roots grow from the nodes. Rhizomes can be long (in couch grass, ground elder) – for rapid dispersal, or short (in iris, Solomon’s seal) – for clump expansion. Separate sections of rhizome containing buds and roots become independent plants after the connecting bridges die off (Serebryakova et al., 2006; Yakovlev et al., 2008). Rhizomes are characteristic of many perennial herbs, shrubs, and even some trees (bamboos).

By tubers. A tuber is a thickened underground shoot with highly developed storage parenchyma. On the tuber surface are “eyes” – axillary buds arranged in a spiral. New shoots develop from the buds, and adventitious roots form on the lower part of the mother tuber (or from stolons). Classic examples: potato, Jerusalem artichoke. Daughter tubers form from stolons (elongated underground shoots) and become independent plants after the stolons die off (Evert & Eichhorn, 2013).

By bulbs. A bulb is a shortened underground shoot (basal plate) with fleshy scales (modified leaves) that store water and nutrients. Daughter bulbs (“offsets”) form in the axils of the scales. After the mother bulb dies, the offsets give rise to new plants. Bulb reproduction is characteristic of tulips, lilies, onions, garlic, hyacinths (Serebryakova et al., 2006).

By stolons (runners). Stolons are above-ground creeping shoots with long internodes. At the nodes of the stolon, leaf rosettes and adventitious roots form. Upon contact with moist soil, the rosettes root, the internodes die off, and the young plants become independent. Typical examples: strawberry (garden and wild), silverweed, some species of spider plant (Bell, 1991; Tan et al., 2025).

By brood buds (bulbils, tubercles, aerial bulbils). Some plants form specialized vegetative reproductive organs directly on the shoots. For example, in viviparous knotweed (Polygonum viviparum), small bulbils form in the inflorescences instead of flowers; in bryophyllum (Kalanchoë daigremontiana), entire plantlets with rudimentary roots develop on the leaf margins, which fall off and root (Garcès et al., 2007). In tiger lily, aerial bulbils form in the leaf axils. In arrowhead, winter buds (turions) form on underwater shoots, giving rise to new plants in spring (Serebryakova et al., 2006; Yakovlev et al., 2008).

Reproduction by roots

By root suckers. In a number of plants (aspen, poplar, cherry, lilac, field bindweed, sow thistle), adventitious buds form on lateral roots. From these buds, above-ground shoots – root suckers – develop, which form their own adventitious roots and after some time separate from the mother plant. Root suckers can arise at a considerable distance from the mother plant, facilitating rapid dispersal (Serebryakova et al., 2006). Some weed plants (sow thistle, creeping thistle) successfully reproduce by root fragments during soil cultivation.

By root tubers. In dahlia, lesser celandine, sweet potato, storage roots (root tubers) bear renewal buds located in the basal part (at the root crown). When a tuber is planted, new shoots and adventitious roots develop from these buds. However, root tubers of most species do not have buds over their entire surface, so a root section with a part of the stem is required for propagation (Bell, 1991; Yakovlev et al., 2008).

Reproduction by leaves

Tiny plantlets with roots visible on the leaf margin of kalanchoe — ©Marshman commons.wikimedia.org CC BY-SA 3.0
Leaf cutting propagation in kalanchoe (Kalanchoë pinnata)
On the leaf of kalanchoe (Kalanchoë pinnata), tiny plantlets with roots form along the leaf blade margin. This is a clear example of natural and artificial vegetative propagation using leaves.

By leaf cuttings (in nature). In some plants (bryophyllum, African violet, some begonias), adventitious buds and adventitious roots form on leaves or their parts (in veins, on the blade margin). In natural conditions, a leaf that falls on moist soil can root and give rise to a new plant. This method is especially characteristic of tropical and subtropical species growing in conditions of high humidity (Bell, 1991; Garcès et al., 2007).

Reproduction by brood fronds and other specialized structures

In some fern species (e.g., Asplenium bulbiferum), brood buds form on the leaves, giving rise to new plantlets. In floating pondweed and other aquatic plants, winter buds (turions) form, which overwinter at the bottom and germinate into new individuals in spring (Serebryakova et al., 2006).

3.2. Artificial vegetative reproduction

Humans have used vegetative reproduction since ancient times to rapidly obtain large quantities of genetically uniform planting material. The main methods of artificial vegetative reproduction are listed below (Hartmann et al., 2011; Carmel et al., 2024).

Reproduction by shoot parts

By stem cuttings. A cutting is a detached part of a shoot (stem with leaves and buds). Cuttings can be:

Green (summer, herbaceous) – cut from current season’s growth (currant, gooseberry, phlox, chrysanthemum). Rooted in a humid environment (in cold frames, under plastic).
Semi-hardwood – cut from plants with partially lignified shoots (lilac, mock orange, rhododendrons).
Hardwood (winter) – taken during the dormant period from lignified annual shoots (grape, currant, willow, poplar, quince). Rooted in spring in open ground or in cold frames (Carmel et al., 2024).
Coniferous (with retained needles) – require a special regime of high humidity and often treatment with rooting stimulants.

By layering. A layer is a part of a shoot that is rooted without being separated from the mother plant. After rooting, the layer is separated and planted as an independent plant. Several types of layering are distinguished:

Horizontal – the shoot is bent to the ground, fastened, and covered with soil in several places (gooseberry, currant).
Serpentine – the shoot is bent and covered with soil only in the middle part (viburnum, honeysuckle).
Vertical – the bush is hilled up with moist soil, and vertical shoots form from the lower buds, which are separated after one year (raspberry, cherry).
Air – a ring of bark is removed from the shoot, wrapped with moist sphagnum moss and plastic film until roots form, then separated (for difficult-to-root plants – lemon, fig, magnolia) (Bell, 1991).

By division of the bush. The plant is dug up and cut into several parts so that each division has roots and renewal buds. Used for perennial herbs (peonies, irises, chrysanthemums, delphinium) and shrubs (currant, barberry, spirea) (Hartmann et al., 2011).

Reproduction by roots

By root cuttings. Sections of roots 5–15 cm long are used, collected in autumn or spring. Root cuttings are planted in a loose substrate; shoots and adventitious roots develop from adventitious buds. This is how raspberry, blackberry, horseradish, and some ornamental plants (panicled phlox, evening primrose) are propagated (Hartmann et al., 2011; Yakovlev et al., 2008).

Reproduction by leaves

By leaf cuttings. A whole leaf or part of it is placed in a moist substrate. At the sites of injury to large veins, adventitious buds and roots form. The method is applicable for African violet, begonia, sansevieria, gloxinia, and many succulents (echeveria, jade plant) (Bidabadi & Jain, 2020; Bell, 1991).

Grafting

Grafting is the artificial joining of parts of different plants (or the same plant) to obtain a single organism. The part that is grafted (cutting or bud) is called the scion. The plant onto which the graft is made is called the rootstock. Grafting allows:

propagation of varieties that root poorly from cuttings;
increased winter hardiness, drought tolerance, resistance to pests and diseases due to the properties of the rootstock;
earlier fruiting;
obtaining dwarf or semi-dwarf forms.

Main grafting methods: whip grafting (simple and improved), budding (grafting with a bud – eye), cleft grafting, bark grafting, side grafting, approach grafting. Grafting is widely used in fruit growing (apple, pear, plum, cherry, citrus), viticulture, and ornamental horticulture (roses) (Evert & Eichhorn, 2013; Hartmann et al., 2011).

Micropropagation (tissue culture in vitro)

This is a modern biotechnological method of vegetative reproduction based on the totipotency of plant cells. A small fragment (explant) is placed aseptically onto a nutrient medium in a test tube or flask. On a medium with specific hormones (auxins, cytokinins), multiple shoots (microshoots) develop from the explant. They are separated and rooted, yielding a large number of genetically identical regenerated plants.

Micropropagation allows:

obtaining thousands and millions of plants from one explant in a short time;
sanitizing planting material from viruses and other pathogens (via apical meristems);
propagating species that are difficult to propagate by traditional methods (orchids, some trees, ferns);
conserving rare and endangered species.

Main stages of micropropagation: selection of donor plant → sterilization → establishment in culture → actual micropropagation (multiplication of microshoots) → rooting in vitro or ex vitro → acclimatization to open ground conditions (Bidabadi & Jain, 2020; Hartmann et al., 2011). The method is widely used in nurseries, floriculture, and for producing virus-free potato and fruit crop material.

4. Stages and mechanisms of regeneration

Regeneration during vegetative reproduction is not a single event but a sequential process that includes several stages, starting from the moment of wounding and ending with the formation of a full-fledged individual. Understanding these stages and mechanisms is necessary for controlling cutting rooting, obtaining callus cultures, and micropropagation.

4.1. General scheme of regeneration during vegetative reproduction

Regardless of the method (cutting, tissue culture, leaf propagation), three main stages can be distinguished (Bidabadi & Jain, 2020; De Klerk et al., 1997, cited in Bidabadi & Jain, 2020):

Dedifferentiation stage (preparation). Cells in the wound zone lose specialized features, activate division, and become capable of new development. A meristematic focus or callus forms.
Induction stage (initiation). Under the influence of internal (hormones, polarity) and external (light, temperature, humidity) factors, cells “decide” on the path of development – whether to form roots, shoots, or embryoids. At this stage, morphogenetic gradients are determined.
Realization stage (differentiation and growth). Visible differentiation of root or shoot primordia, their anatomical formation, and subsequent growth occur.

For rooting of stem cuttings, two stages are more often distinguished (Pincelli-Souza et al., 2024; Carmel et al., 2024):

Root initiation (first 2–5 days) – formation of root primordia from cambium cells or other tissues. At this stage, auxins are critically important.
Root elongation and development – actual growth and branching of roots. Here, high concentrations of auxins can inhibit growth, while gibberellins and other hormones play a greater role.

4.2. Early events: wounding and signaling mechanisms

Wounding is the first and most important event that triggers regeneration. Mechanical tissue damage leads to:

Depolarization of cell membranes and generation of electrical signals.
Release of reactive oxygen species (ROS) – hydrogen peroxide, superoxide anion, which act as signaling molecules (Bidabadi & Jain, 2020).
Activation of gene expression – in particular, of WIND1, WIND2, WIND3, WIND4 (WOUND INDUCED DEDIFFERENTIATION). These transcription factors trigger the dedifferentiation program and increase cell sensitivity to auxins (Bidabadi & Jain, 2020; Rymen et al., 2019, cited in Bidabadi & Jain, 2020).
Changes in hormonal balance. In the wound zone, the concentration of auxin (due to activation of YUC genes – YUCCA, encoding key biosynthesis enzymes) and cytokinins sharply increases. The auxin/cytokinin ratio determines the further path of organogenesis (Bidabadi & Jain, 2020; Chen et al., 2016, cited in Bidabadi & Jain, 2020).

Significance of wounding: in natural vegetative reproduction (rooting of stolons, layers), wounding is not an obligatory condition, since roots form at stolon nodes or in zones of contact with soil without prior wounding. However, in artificial cutting, making a cut (wounding) is necessary to trigger regeneration.

4.3. Callus formation

Collage: roots from callus, shoot-forming callus, embryoid regeneration, compact and friable callus — CC BY 4.0
Stages of organogenesis in plant regeneration using tobacco (<span lang="la" class="biological-name">Nicotiana rustica</span>) as an example
Stages include callus formation, adventitious root and shoot initiation, illustrating dedifferentiation and redifferentiation of cells.

Callus is a mass of unorganized, actively dividing parenchyma cells arising on the wound surface and in deeper tissues. Callus can form:

Directly from cells adjacent to the wound (pericycle, cambium, phloem parenchyma).
From dedifferentiated cells in deeper tissues (medullary rays, cortex parenchyma).

Callus can be friable (easily separated into cells, used for obtaining suspension cultures) or compact (organized). Polarity is preserved in callus, and when transferred to a medium with a specific hormonal composition, it differentiates into adventitious roots (at high auxin content), shoots (at high cytokinin content), or somatic embryoids (Bidabadi & Jain, 2020).

Role of callus in vegetative reproduction:

In direct organogenesis (e.g., rooting of stem cuttings, callus may not form, or it forms in minimal amounts; roots arise directly from cambium and pericycle).
In indirect organogenesis (characteristic of many herbaceous plants and tissue cultures), callus formation is obligatory; organs then differentiate from the callus.
In somatic embryogenesis, embryoids form from callus, which then germinate into plants (Bidabadi & Jain, 2020).

4.4. Dedifferentiation and redifferentiation

Dedifferentiation is the loss by a cell of specialized features (e.g., a parenchyma cell loses its large vacuole, shape, and organelle composition) and the acquisition of the ability to divide. Dedifferentiated cells become similar to meristematic cells: they are small, with dense cytoplasm, a large nucleus, and numerous ribosomes. Dedifferentiation is a reversible process; callus cells can redifferentiate when conditions change (Bidabadi & Jain, 2020).

Redifferentiation is the reverse process: callus cells or dedifferentiated cells in the wound zone reacquire specialized features, forming root, stem, or leaf tissues. Redifferentiation leads to the appearance of adventitious roots, axillary shoots, or somatic embryoids.

Transdifferentiation is a special case of redifferentiation where one type of differentiated cell directly transforms into another (e.g., leaf mesophyll cells into root cortex cells) without an intermediate callus stage. Transdifferentiation is rarely observed in higher plant vegetative reproduction, more common in algae and mosses (Bidabadi & Jain, 2020).

4.5. Adventitious root formation (adventitious rhizogenesis)

Adventitious root formation is a key process in cutting and layering propagation. It goes through several stages:

Cell dedifferentiation – in pericycle, cambium, phloem parenchyma, sometimes in medullary rays. Cells lose vacuoles, nuclei enlarge, proteins and RNA are synthesized.
Initiation of root primordia – active mitotic divisions begin in dedifferentiated cells, forming a small growing point (meristematic focus). Auxins (IAA, IBA, NAA) stimulate this process by activating the expression of WOX11, WOX12, LBD16, LBD29 genes (Bidabadi & Jain, 2020; Liu et al., 2014, cited in Bidabadi & Jain, 2020).
Organization of the root meristem – the typical structure of the root apical meristem with root cap, protoderm, ground meristem, and procambium is formed.
Differentiation of root tissues – primary cortex, central cylinder, vessels, phloem are formed.
Root emergence – roots grow through the stem cortex and emerge on the surface.
Elongation and branching – roots grow in length and form lateral roots (Pincelli-Souza et al., 2024).

Role of auxins: auxins are necessary at the stages of initiation and primordium formation. However, after root emergence, high auxin concentrations suppress their growth (Pincelli-Souza et al., 2024). Therefore, when treating cuttings with rooting stimulants, it is important to observe the concentration and exposure time.

4.6. Adventitious shoot formation (adventitious caulogenesis)

Adventitious shoots can form on root cuttings (root suckers), on leaves (bryophyllum, begonia), or from callus (in tissue culture). The process includes:

Activation of divisions in the pericycle or other tissues → formation of a meristematic focus.
Formation of the shoot apical meristem – which differs from root meristem formation: the shoot meristem has several layers (tunica and corpus) and leaf primordia.
Differentiation of leaves and stem.

To induce shoots, a high level of cytokinins (e.g., kinetin, BAP) and a low level of auxins are required. In tissue culture, a cytokinin/auxin ratio > 1 is used for caulogenesis (Bidabadi & Jain, 2020).

4.7. Polarity during regeneration

The phenomenon of polarity is preserved at all stages of regeneration (see section 2). Even in isolated tissue pieces or callus, auxin moves strictly basipetally (from the apical end to the basal end). This determines that roots arise only at the basal pole. In micropropagation and somatic embryogenesis, polarity is manifested in the fact that embryoids form with a clearly expressed apical-basal gradient, even if they develop from isolated cells (Evert & Eichhorn, 2013; Bell, 1991).

4.8. Programmed cell death and regeneration

During regeneration, some cells in the wound zone die by programmed cell death (PCD). This is not random necrosis but a regulated process that:

Removes damaged cells that could be a source of infection.
Creates space for the growth of new tissues.
Releases signaling molecules (oligosaccharides, peptides) that stimulate division of neighboring cells (Bidabadi & Jain, 2020; Heyman et al., 2016, cited in Bidabadi & Jain, 2020).

It has been established that the transcription factors SOG1 (SUPPRESSOR OF GAMMA RESPONSE 1) and ERF115 (ETHYLENE RESPONSE FACTOR 115) are involved in regulating PCD during regeneration. The mechanisms of PCD in vegetative reproduction are being actively studied, but it is already clear that controlled cell death is an integral part of successful regeneration (Bidabadi & Jain, 2020).

4.9. Stages depending on the propagation method

Different methods of vegetative reproduction have their own characteristics in the sequence of stages:

Rooting of stem cuttings: wounding → callus formation (not obligatory) → initiation of root primordia (cambium, pericycle) → root growth → awakening of axillary buds.
Propagation by root cuttings: wounding → callus (often at both ends) → formation of adventitious buds from callus or directly from pericycle → shoot development → formation of own roots.
Propagation by leaf cuttings: wounding (cutting large veins) → callus from mesophyll and veins → formation of adventitious buds and adventitious roots (often simultaneously) → plantlet formation.
Micropropagation: explant → callus (or without callus) → induction of multiple shoots on cytokinin medium → separation of microshoots → rooting on auxin medium → acclimatization (Bidabadi & Jain, 2020; Hartmann et al., 2011).

5. Factors and conditions for vegetative reproduction

The success of vegetative reproduction – rooting of cuttings, survival of layers, plant yield in micropropagation – depends on a set of factors that can be divided into external (exogenous) and internal (endogenous). Conscious management of these factors can significantly increase propagation efficiency and produce high-quality planting material.

5.1. External (exogenous) factors

External factors include environmental conditions in which rooting or regeneration occurs, as well as the composition of the substrate and nutrient media.

Temperature

Temperature affects the rate of cell division, enzyme activity, hormonal balance, and assimilate transport.

Optimal temperatures for rooting of most temperate-zone cuttings lie in the range of +20…+25 °C. At lower temperatures, processes are slow, and the risk of rot increases. At higher temperatures, transpiration increases, and cuttings may wilt before roots form (Carmel et al., 2024).
Separate temperature regime (bottom heat) is a very effective technique for rooting difficult-to-root species. The substrate temperature is maintained at +22…+25 °C, while air temperature is kept at +15…+18 °C. Such a difference stimulates root development and inhibits premature bud break (Hartmann et al., 2011).
Effect on polarity – at high temperatures (around +30 °C) in some species (e.g., tomato), the polarity of auxin transport is disrupted, which can lead to root formation at the apical end of the cutting (Evert & Eichhorn, 2013).
For in vitro cultures, the optimal temperature is usually +24…+26 °C for most species, but for cold-hardy plants (some conifers) it can be lowered to +18…+20 °C (Bidabadi & Jain, 2020).

Air and substrate humidity

Maintaining high air humidity is a critical condition for rooting cuttings, especially green ones.

Reasons: the cutting lacks roots and cannot absorb water, but it continues to lose water through leaves (transpiration). At low air humidity, the cutting quickly loses turgor, wilts, and dies.
Optimal air humidity for rooting most cuttings is 85–95%. In industrial cutting propagation, it is maintained using misting systems (fine-dispersion spraying) or covering with plastic/glass (cold frames, propagators) (Hartmann et al., 2011).
Substrate humidity should be high but not excessive. Water stagnation causes oxygen deficiency, tissue rot, and cutting death. The substrate should be water-retentive but well-drained (Carmel et al., 2024).
For in vitro cultures, humidity inside the culture vessel is close to 100%, which is necessary to prevent explant desiccation. However, high humidity promotes hyperhydricity (vitrification) – a physiological disorder where tissues become watery and brittle. To reduce hyperhydricity, aeration is increased (Bidabadi & Jain, 2020).

Light

Light has a complex and ambiguous effect on regeneration.

Green cuttings should receive diffused light. Direct sunlight causes overheating and severe desiccation. At the same time, complete darkness reduces photosynthesis, and the cutting consumes carbohydrate reserves, slowing root formation. Optimal light intensity for green cuttings is about 5–10 thousand lux (Hartmann et al., 2011).

Hardwood cuttings (grape, currant, willow) root in darkness or at very low light intensity, as they have no leaves and photosynthesis does not occur. Reserve substances are already accumulated in the bark and wood.

Spectral composition of light. Studies with light-emitting diodes (LEDs) have shown that:

Red light (660 nm) stimulates shoot growth and internode elongation.
Blue light (450 nm) promotes stem thickening, chloroplast development, and adventitious root initiation (Bidabadi & Jain, 2020; Azmi et al., 2014, cited in Bidabadi & Jain, 2020).
Red to blue light ratio (2:1 or 3:1) often gives the best results for in vitro micropropagation.
Etiolation (darkening) of donor shoots is an agrotechnical technique that increases the rooting ability of cuttings. The shoots are wrapped with black paper or foil for several weeks before cutting. In the darkened zone, tissues become looser, mechanical strength decreases, and root emergence is facilitated. A classic example is etiolated grape cuttings, which root better than green ones (Pincelli-Souza et al., 2024).

Substrate and mineral nutrition

The rooting substrate must meet several requirements:

High water-holding capacity – retain water needed by the cutting until roots form.
Good aeration – provide oxygen access to the root formation zone.
Sterility or low contamination – prevent development of fungal and bacterial rots.
Neutral or slightly acidic reaction (pH 5.5–6.5) – for most species.

Typical substrates for cutting propagation: peat and sand mixture (1:1), perlite, vermiculite, coconut coir, and combinations thereof. For difficult-to-root species, clean washed sand or perlite is used to minimize the risk of infection (Carmel et al., 2024).

Mineral nutrition during the rooting period should be balanced:

High phosphorus content stimulates root system development.
Potassium increases resistance to stress and water deficit.
Nitrogen should be limited at the initial rooting stage, as excess nitrogen stimulates shoot growth at the expense of roots (Hartmann et al., 2011).
Industrial substrates often contain starter fertilizer doses with low nitrogen and elevated phosphorus and potassium (e.g., 5-20-10 or 10-20-10).

Phytohormones and growth regulators

Regulation of vegetative reproduction using exogenous phytohormones is one of the most powerful and widely used techniques. Main groups:

Auxins (indole-3-acetic acid – IAA, indole-3-butyric acid – IBA, 1-naphthaleneacetic acid – NAA).
- IBA is the most effective rooting stimulant for most species. Unlike IAA, IBA is more stable to light and oxidation, and in cutting tissues it is converted into active IAA, creating a long-lasting gradient (Pincelli-Souza et al., 2024).
- NAA is a synthetic auxin, often used in mixture with IBA to enhance the effect.
- Concentrations: from 500 to 5000 mg/L (and higher) for quick “dry” or “liquid” treatment of the cutting base (dip method). For aqueous solutions – 25–100 mg/L with soaking for 12–24 hours (Carmel et al., 2024).
- Auxins not only accelerate root emergence but also increase the number of roots per cutting.
Cytokinins (kinetin, benzylaminopurine – BAP, zeatin).
- Cytokinins stimulate the formation of adventitious buds and shoots. They are used in propagation by root cuttings (raspberry, phlox) and in tissue culture.
- Excess cytokinins suppresses rooting, so they are rarely used at the rooting stage of stem cuttings.
- The auxin/cytokinin ratio regulates the path of morphogenesis: >1 → roots, <1 → shoots (Bidabadi & Jain, 2020).
Gibberellins (GA3 and others).
- Usually suppress root formation, stimulating shoot growth. However, at low concentrations they may enhance rhizogenesis in some species (Carmel et al., 2024).
- Used to stimulate growth of already rooted cuttings.
Inhibitors and other regulators. In some cases, gibberellin inhibitors (paclobutrazol, ancymidol) are used to improve rooting, especially in woody species. Phenolic compounds (phloroglucinol, rutin) are also used as auxin synergists (Hartmann et al., 2011).
Current trends: use of nanoparticles (nanosilver, metal oxide nanoparticles) and new synthetic compounds (e.g., sustained-release IBA derivatives) to stimulate rooting (Bidabadi & Jain, 2020; Pincelli-Souza et al., 2024).

5.2. Internal (endogenous) factors

Internal factors are related to the donor plant itself and the physiological state of the detached part.

Age of the donor plant

Juvenile (young) plants root significantly better than adult ones. This is due to higher auxin content, less lignification of tissues, and greater hormonal plasticity.

Aging effect in vegetative reproduction has been well studied in woody species: cuttings taken from adult trees root poorly, and plants derived from them take a long time to flower.

Rejuvenation – a set of techniques (grafting onto a juvenile rootstock, repeated cuttings, apical meristem culture) that return an adult plant to a state close to juvenile in terms of regeneration ability (Hartmann et al., 2011). Widely used in forestry and horticulture.

Type and quality of cutting (position on the shoot, length, diameter)

Position on the shoot: cuttings from the middle part of the shoot often root better than those from the upper or lower part. In many species, basal cuttings contain more storage reserves but may be more lignified (Carmel et al., 2024).

Cutting length affects the supply of nutrients and the transpiration area. Optimal length for most species is 10–15 cm for green cuttings and 15–25 cm for hardwood cuttings.

Diameter correlates with carbohydrate reserves and auxin content. Too thin cuttings exhaust quickly, too thick ones form roots slowly.

Presence of leaves on green cuttings is mandatory (except for some species). Leaves are a source of auxins and carbohydrates. Leaf removal sharply reduces rooting ability.

Physiological state of the donor and seasonality

Season of cutting collection is one of the most important factors.

Hardwood cuttings (fruit, grape, currant) are collected during the dormant period (autumn after leaf fall or early spring before bud swell). Stored at –1…–2 °C in moist sawdust or sand.

Green cuttings are cut during the active shoot growth period (late May – July). Time of day – early morning, when tissues are maximally saturated with water.

Semi-hardwood cuttings are collected in July – August, when the shoot base begins to lignify but the tip is still growing.

Effect of season on rooting ability is related to changes in endogenous levels of hormones, carbohydrates, and inhibitors. For example, in apple cuttings taken in winter, rooting is minimal, while in spring it is maximal (Carmel et al., 2024).

Donor plant condition. Healthy, well-lit, well-watered, and well-fed plants produce cuttings with high rooting energy. Lack of light, excess nitrogen, or drought reduce cutting quality.

Hereditary (genotypic) characteristics

Species and varietal specificity – some species root easily (willow, poplar, currant, grape), others are difficult or do not root at all (pine, oak, walnut, most fruit trees on their own roots).

Rooting ability is a heritable polygenic trait. Selection for improved cutting rooting is possible, and such varieties exist (e.g., some apple rootstocks).

Genetic mechanisms include differences in expression of auxin biosynthesis genes (YUC), auxin receptor genes (TIR1/AFB), and transcription factors (WOX11, WOX12, LBD16/29) (Bidabadi & Jain, 2020; Pincelli-Souza et al., 2024). In difficult-to-root genotypes, the ability to conjugate auxins is often reduced, or their transport is impaired.

Nutrient reserves

Carbohydrates (starch, sugars) are the main energy and building material for root formation. Cuttings with high starch content root better. In hardwood cuttings, carbohydrate reserves in bark and wood are critically important.

Nitrogen compounds and other elements are also necessary. However, excess nitrogen can shift the balance toward vegetative growth at the expense of rooting (Hartmann et al., 2011).

Phenolic compounds – some of them (e.g., phloroglucinol) act as auxin synergists, protecting IAA from oxidation. In difficult-to-root species, the content of endogenous phenols is often lower.

5.3. Interaction of factors

In real conditions, factors do not act in isolation but in complex interaction. For example, the effectiveness of auxin treatment depends on temperature, humidity, donor age, and cutting type. For each species and variety, an optimal protocol is developed, taking into account:

cutting collection time;
cutting type (green, semi-hardwood, hardwood);
concentration and composition of rooting stimulants;
substrate and irrigation regime;
temperature and light regime.

Only a systemic approach allows achieving maximum rooting percentages and yield of high-quality planting material.

6. Management techniques and applied significance

Management of vegetative reproduction is a set of agrotechnical and biotechnological techniques aimed at stimulating regeneration processes (primarily rooting), increasing the yield of standard planting material, and accelerating the propagation of valuable genotypes. These techniques have enormous practical importance for agriculture, horticulture, forestry, ornamental plant production, and biotechnology.

6.1. Techniques for managing vegetative reproduction

Stimulation of root formation (rhizogenesis)

The main problem in cutting propagation of many species (especially woody and difficult-to-root ones) is the weak ability to form adventitious roots. The following techniques have been developed to overcome it.

Use of synthetic auxins

Auxins are the most effective stimulators of rhizogenesis. In practice, the following are used:

Indole-3-butyric acid (IBA). Considered the “gold standard” for cutting propagation. IBA is more stable to light and oxidation than IAA, and is slowly metabolized in tissues, creating a long-lasting gradient (Pincelli-Souza et al., 2024). Available as salts (potassium salt of IBA – for aqueous solutions), powders (with talc), and gels.
1-Naphthaleneacetic acid (NAA). Often used in mixture with IBA (e.g., 2:1 or 1:1). NAA is particularly effective for some woody species (Carmel et al., 2024).
Indole-3-acetic acid (IAA). Natural auxin. Less stable than IBA, but also used, especially in combinations.
New synthetic compounds: halogenated IAA derivatives (4-Cl-IAA, 5,6-diCl-IAA) have higher activity and selectivity; esters and auxin conjugates with prolonged action are also being studied (Pincelli-Souza et al., 2024).

Methods of treating cuttings with auxins (Hartmann et al., 2011; Carmel et al., 2024):

Dry dip method (powder dipping). The base of the cutting (1–2 cm) is moistened with water and dipped into a powder containing talc and auxin (usually 0.5–2% IBA). The most common method in amateur and industrial horticulture.
Liquid dip method (quick dip). The base of the cutting is dipped into a high-concentration alcohol or alcohol-water auxin solution (500–5000 mg/L) for 3–5 seconds. Economical for mass treatment.
Prolonged treatment in aqueous solution. Cuttings are soaked in a weak auxin solution (25–100 mg/L) for 12–24 hours. Effective for species with difficult-to-root cuttings.
Gel or paste treatment. Convenient for vertical cuttings and grafting.

Effectiveness of treatment depends on the plant species, concentration, and exposure time. For some species, very high concentrations (2000–4000 mg/L and above) are optimal; for others, medium concentrations (500–1000 mg/L) (Carmel et al., 2024; Pincelli-Souza et al., 2024).

Other regulators and synergists

Phenolic compounds (phloroglucinol, rutin, catechol) – enhance the action of auxins, protecting them from oxidation. Sometimes added to auxin solutions.
Gibberellin inhibitors (paclobutrazol, uniconazole) – can improve rooting in some woody species by blocking gibberellin synthesis, which inhibits rhizogenesis (Pincelli-Souza et al., 2024).
Auxin conjugation inhibitors (keikimid, nalacin) – block GH3 enzymes that conjugate auxin with amino acids, thereby increasing free auxin levels. In experimental research stage (Pincelli-Souza et al., 2024).

Agrotechnical techniques for preparing donor plants and cuttings

Etiolation (darkening) of shoots. Shoots are wrapped with opaque paper or foil 2–4 weeks before cutting collection. In the darkened zone, tissues become looser, mechanical strength decreases, and ethylene- and auxin-induced root primordia formation is activated. Widely used for grapes, roses, and some fruit species (Hartmann et al., 2011).

Kilching (stratification of cuttings). Cuttings (especially hardwood) before planting are placed in a moist substrate at elevated temperature (+18…+25 °C) to accelerate callus formation and root initiation. Used for grape, actinidia, and schisandra cuttings.

Scarification and bark scoring. Shallow longitudinal scratches or narrow strips of bark are removed from the lower end of the cutting. This increases the contact area with the substrate and facilitates root emergence.

Disinfestation (sterilization) of substrate and cuttings. To prevent fungal diseases (damping-off, fusarium), the substrate is steamed or treated with fungicides, and cuttings are treated with fungicide solutions or biological agents.

Nitrogen and phosphorus treatment. Donor plants are fed with phosphorus-potassium fertilizers 2–3 weeks before cutting collection. Excess nitrogen reduces rooting.

Techniques in tissue culture (micropropagation)

Nutrient medium selection. Composition of macro- and microelements, vitamins, carbohydrates (usually 2–3% sucrose). At the shoot multiplication stage, a high cytokinin/auxin ratio is used; at the rooting stage, a low ratio (Bidabadi & Jain, 2020).

Light culture. Use of LEDs with adjustable spectrum (red+blue) increases the yield and quality of microshoots.

Photoautotrophic micropropagation. Growing on medium without sucrose, at elevated CO₂ concentration (1000–1500 ppm) and high light intensity. Reduces hyperhydricity and lowers cost (Bidabadi & Jain, 2020).

Liquid cultures and bioreactors. Suspension cultures and temporary immersion systems (TIS) allow process automation and scaling up production.

Sanitation through meristem culture. Isolating apical meristems (0.1–0.5 mm in size) produces virus-free planting material, as viruses usually do not penetrate apical meristems (Sakthivel et al., 2025).

Grafting: managing cultivar and adaptive traits

Grafting not only propagates a cultivar but also imparts new properties to the plant through the rootstock:

Growth vigor regulation – dwarf, semi-dwarf, and vigorous rootstocks (for apple, pear).
Increased winter hardiness, drought tolerance, salt tolerance – through the resistant rootstock.
Resistance to pests and diseases (e.g., apple rootstocks resistant to root rot and woolly apple aphid).
Earlier fruiting (many fruit trees on dwarf rootstocks begin fruiting in the 2nd–3rd year).
Overcoming incompatibility – interstock (intermediate grafting).

Main grafting methods: improved whip grafting, budding (eye grafting), cleft grafting, bark grafting, approach grafting. Choice of method depends on the time of year (spring – whip grafting, summer – budding), thickness of rootstock and scion (Hartmann et al., 2011).

Modern molecular approaches

Use of regeneration ability biomarkers. Expression levels of WOX11, WOX12, LBD16, LBD29 genes in cutting tissues can serve as predictors of rooting ability (Bidabadi & Jain, 2020).

Genome editing to enhance vegetative reproduction ability. For example, editing GH3 (auxin conjugation) or DMR6 (cultivar resistance) genes in difficult-to-root crops (Sakthivel et al., 2025).

Breeding clones with improved regeneration ability. Use of marker-assisted selection (MAS) and genomic selection to create varieties and rootstocks that are easily propagated vegetatively (Sakthivel et al., 2025).

6.2. Applied significance of vegetative reproduction

Vegetative reproduction is the foundation of modern intensive plant cultivation. Its importance can hardly be overstated.

Agriculture and horticulture

Propagation of fruit and berry cultivars. Almost all varieties of apple, pear, plum, cherry, currant, gooseberry, raspberry, strawberry are propagated vegetatively (by grafting, cuttings, runners, division of the bush). This guarantees preservation of cultivar traits (Hartmann et al., 2011).
Vegetable growing. Potato (tubers), sweet potato (cuttings and tubers), Jerusalem artichoke (tubers), horseradish (root cuttings), onion and garlic (bulbs and cloves), mint and other herbs (cuttings and division).
Viticulture. Grape is propagated mainly by hardwood and green cuttings, as well as by grafting onto phylloxera-resistant rootstocks.
Nursery production. Production of planting material for fruit, berry, ornamental, and forest crops on an industrial scale (millions of seedlings per year) is possible only through vegetative reproduction.

Forestry

Propagation of plus trees – individuals with superior growth, trunk quality, and resistance in forest species breeding programs (pine, spruce, larch, poplar). Cutting and micropropagation allow multiplying elite genotypes (Sakthivel et al., 2025).
Rapid propagation of fast-growing species (poplar, willow, eucalyptus) for plantation forestry and biofuel.
Conservation of rare and endangered tree species through tissue culture and clonal collections.

Ornamental horticulture and floriculture

Propagation of roses (cuttings, grafting).
Propagation of conifers (thuja, cypress, juniper, yew) – by hardwood and green cuttings, often using misting systems.
Propagation of perennial flowers (phlox, chrysanthemum, lily, iris, peony) by division of the bush, cuttings, bulbs.
Indoor floriculture – African violets, begonias, ficuses, ivies, many succulents are easily propagated by leaf and stem cuttings.
Obtaining new colors and forms (chimeras) through vegetative propagation of sports (bud mutations).

Biotechnology and fundamental research

Infographic: potato, banana, strawberry, sugarcane; stresses, diseases, quality; transgenesis and CRISPR/Cas9 — CC BY 4.0
Application of transgenic and genome-edited technologies for improving vegetatively propagated polyploids
Modern biotechnological methods allow creating clones with desired traits while preserving cultivar characteristics.

Micropropagation allows obtaining tens and hundreds of thousands of plants from one explant in a short time. This is especially important for crops where traditional propagation is difficult (orchids, ferns, pineapples, many woody species) (Bidabadi & Jain, 2020).
Sanitation of planting material. Meristem culture is the main method for obtaining virus-free potato, strawberry, fruit, citrus, and ornamental crops.
In vitro germplasm conservation. Germplasm of rare and economically valuable species is stored in tissue culture collections at low temperatures (cryopreservation) or under slow growth.
Genetic engineering and genome editing. Plant transformation and gene editing are possible only through regeneration from somatic cells (callus, protoplasts) – i.e., through vegetative reproduction in vitro. Obtaining transgenic and genome-edited plants requires optimization of regeneration protocols for each genotype (Sakthivel et al., 2025).
Study of morphogenesis, differentiation, and polarity mechanisms. Vegetative reproduction is a convenient model for fundamental research in plant developmental biology.

Agricultural economics and food security

Reducing the time from cultivar release to industrial production (from 10–15 years with seed propagation to 2–4 years with vegetative propagation).
Ensuring product uniformity (in size, taste, ripening time) – critical for mechanized harvesting and processing.
Rapid propagation of new resistant varieties in response to the emergence of new pathogen races (e.g., late blight‑resistant potato varieties, scab‑resistant apple).
Saving space for planting material production (in micropropagation).

6.3. Successful application examples

Spider plant (Chlorophytum comosum) – produces numerous rosettes on stolons that root easily; one of the most common houseplants propagated vegetatively on an industrial scale.
Potato – all world seed potato is propagated vegetatively (by tubers or microtuberlets), which preserves cultivar traits but creates a virus accumulation problem, solved by meristem sanitation.
Apple – all commercial varieties are propagated by grafting onto clonal rootstocks (M9, MM106, 54–118, etc.). Without vegetative reproduction, modern intensive orcharding would be impossible.
Orchids – many species (Phalaenopsis, Cymbidium, Dendrobium) are propagated exclusively in vitro via protocorms – embryo-like structures developing from seeds or meristems. Only micropropagation has made orchids mass‑market houseplants (Hartmann et al., 2011; Bidabadi & Jain, 2020).
Poplar and willow plantations for biofuel and pulp‑and‑paper industry are established from cuttings (hardwood or green) of the best clones selected for fast growth and wood quality.

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