Virginal (Vegetative) Stage

Last updated: June 12, 2026EspañolРусский

Ontogenesis of a flowering plant, beginning with the fertilized egg cell and ending with natural death, is commonly divided into several major periods. Among these, in addition to the latent (seed) and reproductive (flowering and fruiting) periods, a central place is occupied by the so-called pre-reproductive (or virginal) period — the stage in a plant’s life when it has already formed as a vegetatively mature organism but has not yet begun sexual reproduction.

Virginal (vegetative) stage of ontogenesis (from Latin virgo — maiden, virgin; English virginal stage, adult vegetative phase) — this is the post-embryonic developmental stage of a seed plant during which the individual achieves the vegetative structure typical of the species, exhibits a high intensity of growth processes, but is physiologically incapable of forming generative organs (flowers, fruits, seeds) (Gatsuk et al., 1980; Yakovlev et al., 2008).

In the Russian botanical school, following the classic works of T. A. Rabotnov and A. A. Uranov, the virginal stage is distinguished as one of the age states (cenopopulation states) alongside the immature (transitional) and generative stages (Serebryakova et al., 2006). In foreign literature, the term “adult vegetative phase” often corresponds to this, which is distinguished from the juvenile phase (Zotz et al., 2011).

Key characteristics of the virginal stage:

  1. Vegetative maturity. The plant is no longer a seedling or a juvenile individual. Its leaves, stems, and roots acquire morphological features typical of an adult plant of that species: juvenile leaf forms disappear (heteroblasty), characteristic venation, pubescence, and leaf blade shape develop. For example, in ivy (Hedera helix) the palmate-lobed juvenile leaves are replaced by entire, rhombic adult leaves of the virginal stage (Zotz et al., 2011).

  2. Intensive growth. Unlike the subsequent generative period, when part of the resources are allocated to flowering and fruiting, biomass increment is maximal during the virginal stage. This applies to both above-ground (shoot elongation, increase in leaf number) and underground organs (root system development).

  3. Absence of generative organs. The decisive diagnostic characteristic. The plant produces neither flowers nor fruits, although its growing point (apex) may already be prepared for transition to a reproductive state, but this transition is inhibited by endogenous or exogenous factors.

  4. Capacity for vegetative reproduction. During the virginal stage, many species actively reproduce vegetatively (by rhizomes, layering, stolons), contributing to the formation of clones and thickets. This distinguishes virginal individuals from juvenile ones, which are generally not yet capable of vegetative renewal (Gatsuk et al., 1980).

Differences from adjacent age states. The virginal stage occupies a strictly defined place in the ontogenetic sequence:

  • After the immature stage (sub-adult individuals), where individual juvenile features are still retained, and the plant is only beginning to acquire adult structural characteristics.

  • Before the generative stage, when the first flower buds are initiated.

Thus, the virginal state is precisely that “adult, but not yet flowering” period which, for many cultivated plants (cabbage, beet, carrot), holds the greatest economic value — for it is during this time that the main vegetative mass is formed (Yakovlev et al., 2008).

Variability in duration and transition factors. The duration of the virginal stage varies widely depending on taxonomic affiliation, life form, and environmental conditions. In annual ephemerals, it may last days; in long-lived trees (oak, beech) — decades (Gatsuk et al., 1980). The key factors determining the moment of exit from the virginal state are photoperiod, temperature regime (vernalization), and the plant’s hormonal status. The transition to flowering itself is often not rigidly linked to reaching a specific chronological age — the same individual, under different environmental conditions, may remain virginal significantly longer or, conversely, flower earlier.

Thus, the virginal stage of ontogenesis is not merely a “pause” before flowering, but a crucial period of active biomass accumulation and structural organization of the plant, largely determining its future productivity and reproductive success.

1. Morphological and structural changes during the transition to the virginal state

The transition of a plant from the juvenile (or immature) state to the virginal state never occurs instantaneously. It is a gradual but fairly distinct process affecting all vegetative organs. Externally, it manifests in changes in leaf shape, branching pattern, stem structure, and root system. Internally, it involves the complexification of anatomical structure, the appearance of secondary tissues, and the completion of conducting system formation. Let us examine the key changes.

1.1. Cessation of heteroblasty and establishment of the “adult” leaf form

Tree with juvenile foliage at the base and adult foliage at the top

Juvenile and adult foliage of the tree <span lang="la" class="biological-name">Elaeocarpus hookerianus</span>

Photograph of a tree clearly showing the coexistence of different leaf types: juvenile (at the base) and adult, typical of the virginal stage (in the crown).

One of the most noticeable events during the transition to the virginal stage is the disappearance of heteroblasty — the phenomenon where leaves of different shapes are formed on the same plant at different ages (Zotz et al., 2011). In many species, juvenile leaves (the first leaves of the seedling) have a simpler, sometimes lobed or even thread-like form. Upon reaching the virginal state, all newly initiated leaves acquire the shape, size, and venation pattern typical of the adult plant of the species.

Examples:

  • In blackwood acacia (Acacia melanoxylon) juvenile leaves are bipinnate, compound; at the virginal stage, they are replaced by flattened, leaf-like petioles — phyllodes (Zotz et al., 2011).

  • In ivy (Hedera helix) juvenile leaves are palmate-lobed, virginal leaves are entire, ovate-rhombic.

  • In blue gum eucalyptus (Eucalyptus globulus) a transition is observed from sessile, opposite, glaucous juvenile leaves to petiolate, alternate, glossy green adult-type leaves.

From the perspective of morphogenesis, the cessation of heteroblasty indicates the completion of the restructuring of the shoot apical meristem. During this period, leaf primordia are initiated not according to the juvenile pattern, but according to the adult pattern, which is controlled by internal (genetic, hormonal) mechanisms and is generally irreversible (Zotz et al., 2011).

1.2. Changes in phyllotaxis and shoot branching pattern

In many woody and herbaceous plants, the arrangement of leaves on the stem (phyllotaxis) may change during the transition to the virginal stage. For example, in seedlings of Scots pine (Pinus sylvestris), needles are arranged alternately, while in the adult virginal plant they are arranged in fascicles on short shoots (Gatsuk et al., 1980).

Simultaneously, the branching pattern often changes. At the juvenile stage, shoots generally branch weakly (monopodially without significant lateral axis formation). Upon reaching the virginal state, the initiation of lateral shoots is activated, and a skeletal branch system is formed. In trees, this manifests in the appearance of branches of the second and subsequent orders; in herbs, in intensive tillering or the formation of rhizomatous shoots (Serebryakova et al., 2006).

1.3. Changes in stem and root structure

The transition to the virginal state is accompanied not only by external changes but also by profound anatomical changes:

  • Stem thickening. In dicotyledons and gymnosperms, the cambium is activated, and secondary stem thickening begins. Well-developed wood (xylem) and bast (phloem) are formed. This provides mechanical strength and increases the capacity of the conducting pathways.

  • Periderm (cork) development. The epidermis is replaced by a multilayered cork, protecting the stem from desiccation and mechanical damage.

  • Complexification of the root system. The primary root of the seedling (taproot) branches intensively, forming lateral roots of the first, second, and subsequent orders. In monocotyledonous plants, the taproot often dies off and is replaced by adventitious roots growing from stem nodes. A fibrous root system, characteristic of adult virginal grasses, is formed (Gatsuk et al., 1980).

1.4. Change in growth pattern (open growth system)

The virginal plant differs from the juvenile one in its capacity for prolonged, indeterminate growth (until the generative phase arrives). In trees, this is expressed in the annual elongation of the main shoot and branches. In perennial herbs during the virginal stage, perennial underground shoots (rhizomes) are formed, which each year produce new above-ground increments. Thus, the plant vegetatively “rejuvenates,” retaining the ability to colonize new areas (Serebryakova et al., 2006).

Key conclusion: The morphological and structural changes during the transition to the virginal state are all directed towards one goal — creating a powerful, autonomous, and stable vegetative system capable of ensuring maximal possible seed productivity in the future. This is a stage of “investment” in its own skeleton and conducting network.

2. Characteristics of the stage in different life forms

The virginal stage is not universal in its duration and internal structure. It varies significantly depending on the plant’s life form (tree, shrub, perennial or annual herb). These differences have direct practical significance for the agronomist, as they determine the timing of fruiting onset, the duration of use of vegetative mass, and the crop management system.

The classification of life forms is based on the location of renewal buds relative to the soil surface and the degree of shoot lignification (Serebryakova et al., 2006). Let us examine the characteristics of the virginal stage for the main groups.

2.1. Annual herbs (ephemerals and spring crops)

In annual plants, the entire life cycle fits into one growing season. The virginal stage in them is maximally compressed (Gatsuk et al., 1980).

  • Duration: From a few days (in desert ephemerals) to 2–4 weeks (in most spring cereals and row crops).

  • Characteristics: The plant quickly passes through the seedling stage (cotyledon leaves) and the juvenile stage (first true leaves), after which it almost immediately transitions to the formation of “adult” leaves and stem. The virginal phase in annuals often coincides with the “tillering” period (in cereals) or “branching” period (in legumes).

  • Agricultural significance: For annual crops (wheat, barley, corn, sunflower), the virginal stage is critically important for yield formation of vegetative mass. Any stress (drought, nitrogen deficiency) during this period reduces the number of productive tillers and, consequently, the future yield of grain or green mass.

2.2. Biennial plants

Biennials (carrot, beet, cabbage, parsley) represent a particular strategy: the virginal stage occupies almost their entire first year of life (Gatsuk et al., 1980; Yakovlev et al., 2008).

  • Duration: The entire first growing season (sometimes extending into part of the second summer before flowering).

  • Characteristics: During this period, the plant does not flower but accumulates reserve nutrients in specialized modified organs: root vegetables (carrot, beet), heads (cabbage), bulbs. All morphological changes are directed towards creating a storage structure. Leaves form a rosette, the stem remains shortened.

  • Agricultural significance: It is precisely the virginal stage that is “economically useful” for biennials. The yield of root vegetables, heads, and bulbs is harvested before the plants transition to the generative phase. If a biennial flowers in its first year (e.g., due to early planting or after a cold spring — “vernalization”), the commercial quality of the product sharply decreases (the root vegetable becomes “woody,” the head cracks).

2.3. Perennial herbs (polycarpic)

In perennial herbs (clover, alfalfa, timothy, couch grass), the virginal stage can last several years, with annual rhythmic renewal of vegetation (Gatsuk et al., 1980; Serebryakova et al., 2006).

  • Duration: From 1–2 years (in some meadow grasses) to 5–10 years (in leguminous herbs and rhizomatous species).

  • Characteristics: The plant forms a polycentric structure: a perennial rhizome (or caudex) with annually regrowing annual shoots. During the virginal stage, underground organs (rhizomes, stolons) are initiated and expand, leading to vegetative spread of the clump.

  • Agricultural significance: For perennial forage grasses, the virginal stage is the period of greatest productivity of green mass. Mowing the stand during this period (tillering phase — beginning of stem elongation) not only yields a high harvest but also stimulates further vegetative renewal. Transition to the generative phase (flowering) leads to stem coarsening and reduced nutritional value.

2.4. Trees and shrubs

In woody forms, the virginal stage is the longest in the plant kingdom (Gatsuk et al., 1980; Yakovlev et al., 2008).

  • Duration: From 3–5 years (in some fruit shrubs, e.g., currant) to 20–40 years or more (in forest trees: oak, beech, pine).

  • Characteristics: The long period of vegetative growth is necessary for forming a strong skeleton — the trunk and main crown branches. During this time, active secondary thickening (cambium) occurs, and a powerful root system is formed (Gatsuk et al., 1980). Young trees (saplings) in the nursery are precisely in the virginal state. They do not yet bear fruit, but have already formed the crown shape typical of the species.

  • Agronomic significance:

  • Pomology: The duration of the virginal stage is an important breeding trait. Precocious varieties of apple, pear, and cherry have a shortened virginal period (3–5 years) compared to wild forms (10–15 years).

  • Forestry: In forestry, the age of crown closure and the onset of intraspecific competition occurs at the end of the virginal stage.

  • Ornamental horticulture: Formative pruning (creating the trunk, scaffold branches) is carried out precisely during the virginal stage, as the plant is still plastic and does not expend resources on flowering.

2.5. Subshrubs and dwarf shrubs

This life form (wormwood, anabasis, thyme, lingonberry) occupies an intermediate position (Gatsuk et al., 1980).

Characteristics: The virginal stage in them is relatively short (1–2 years). The plant quickly forms lignifying lower parts of shoots and non-lignifying (“herbaceous”) upper growths. A specific feature is the partial dieback of shoots after fruiting. During the virginal stage in subshrubs, the base of the future bush — the basal part with renewal buds — is established (Serebryakova et al., 2006).

3. Stage duration and factors affecting its length

The virginal stage does not have a rigidly fixed calendar duration, even among individuals of the same species. Its length varies widely and is determined by the complex interaction of the genetic program (endogenous factor) and environmental conditions (exogenous factors). Understanding these mechanisms allows the agronomist to consciously manage the plant’s transition from vegetative growth to flowering.

3.1. Genetic determinacy (species specificity)

The fundamental boundary of the possible duration of the virginal stage is embedded in the genotype. Different species, and even varieties of the same species, have a hereditarily fixed “norm” of time required to reach vegetative maturity (Gatsuk et al., 1980; Yakovlev et al., 2008).

  • Monocarpics (annuals): Genetically programmed for an extremely short virginal stage (days, weeks). Their task is to form a minimum of vegetative mass as quickly as possible and transition to flowering.

  • Polycarpic herbs and shrubs: The virginal stage lasts from 1–2 to 5–10 years. For example, in red clover (Trifolium pratense) the virginal state may continue for 2–3 years before the plant flowers (Gatsuk et al., 1980).

  • Trees: The longest virginal period. In European crab apple (Malus sylvestris) it reaches 10–15 years, in pedunculate oak (Quercus robur) — 20–40 years, and in European beech (Fagus sylvatica) — up to 50 years (Gatsuk et al., 1980).

This hereditarily fixed duration is the result of evolutionary adaptation to specific habitat conditions. Species growing in competitive environments (forests) are forced to “invest” many years in height growth and the development of a powerful trunk to gain access to light and successfully reproduce in the future.

3.2. Exogenous factors regulating the stage duration

External conditions can either accelerate or delay the exit from the virginal state. The key factors are:

Light (photoperiodism and intensity)

Photoperiod (day length): For many species, the transition to flowering is triggered only at a specific day length (short-day plants, long-day plants, or day-neutral). While the plant is in the virginal stage, it is either not receptive to photoperiod, or it is receptive but conditions are not yet suitable. However, a change in photoperiod can accelerate the end of the virginal stage. For example, in species of hemp (Cannabis) and chrysanthemum, short days induce flowering, thereby shortening the virginal phase (Mauseth, 2016).

Light intensity (shading): Under light deficiency (in dense crops, under forest canopy), the virginal stage in many species lengthens. The plant “waits” for favorable conditions, elongates upwards, but does not transition to flowering. This phenomenon is especially pronounced in tree species (spruce, fir), which can remain in the virginal state in the understory for decades.

Temperature (vernalization)

For many biennial and perennial plants, exposure to low positive temperatures (vernalization) is an obligatory condition for exiting the virginal stage. Without experiencing a cold period, the plant remains in the virginal state and does not flower (Mauseth, 2016).

Mechanism: Perception of cold (usually by the shoot apex) triggers a metabolic rearrangement and the synthesis of signaling molecules (e.g., FT protein — Florigen), which ultimately initiate flower formation.

Agronomic example: Winter varieties of wheat and rye have been selected to require vernalization. They are sown in autumn, undergo the virginal stage (tillering) before winter, and in spring, after vernalization, quickly transition to stem elongation and flowering. If winter wheat is sown in spring, it will remain in the virginal phase, will not head, and will not produce grain.

Mineral nutrition (especially nitrogen and phosphorus)

Excess nitrogen (especially combined with a deficiency of phosphorus and potassium) prolongs the virginal stage. The plant “luxuriates”: it forms a robust vegetative mass (dark green leaves, thick stems) but delays flowering. This phenomenon is well known in crop production as “luxuriant growth” of cereals or vegetable crops (Yakovlev et al., 2008).

Phosphorus deficiency, conversely, can retard development and lengthen the virginal stage, as phosphorus is necessary for energy metabolism and cell division.

Balanced nutrition (especially NPK) ensures a normal developmental rate and timely completion of the virginal stage.

Water regime and stress

Drought, salinization, low temperatures (above the damage threshold) often act as stress signals, accelerating the transition to flowering. A plant in the virginal stage experiencing stress strives to complete its life cycle as quickly as possible and leave seeds (“stress-induced flowering”). Conversely, excessive moisture (waterlogging) can delay development and prolong the virginal phase.

3.3. Endogenous factors (hormonal regulation)

The internal developmental program is controlled by the balance of hormones.

Gibberellins (GA) and auxins stimulate vegetative growth and maintain the virginal state. * The transition to flowering is often associated with a decrease in gibberellin levels and/or an increase in the levels of florigen (FT protein) and other signaling molecules, which are synthesized in leaves in response to external stimuli (photoperiod, vernalization) and transported to the shoot apex (Mauseth, 2016).

Treatment of plants with exogenous gibberellins can, in some cases, mimic vernalization and accelerate the transition to flowering (e.g., in biennial Brassicaceae plants), thereby shortening the virginal stage.

4. Criteria for the beginning and end of the stage

For accurate determination of the age state of a plant under field conditions (in agrocenoses, forest stands, pastures), it is necessary to rely on clear, easily recognizable diagnostic characteristics. These characteristics allow objective recording of the moment an individual enters the virginal stage and the moment it ends, even without knowing the plant’s chronological age (Gatsuk et al., 1980; Serebryakova et al., 2006).

It is important to emphasize that the transition from one age state to another is a gradual process, but there are critical morphological shifts that serve as reliable markers.

4.1. Criteria for the onset of the virginal stage

An individual is considered to have entered the virginal state (v) when it loses all juvenile characteristics and acquires the structure typical of an adult plant, but has not yet begun to flower (Gatsuk et al., 1980; Yakovlev et al., 2008). The main criteria are:

  1. Cessation of heteroblasty. All newly formed leaves have the shape, venation, pubescence, and color characteristic of the adult plant of the species. Juvenile-type leaves (simple, small, of a different shape) no longer appear (Zotz et al., 2011).

  2. Completion of the formation of the “adult” shoot type. In trees and shrubs: formation of the first full tier of scaffold branches, appearance of the crown typical of the species. In herbs: completion of tillering and transition to a hollow stem (in grasses) or the formation of a developed rosette with leaves of the median formation.

  3. Formed root system. In dicotyledons: a distinct taproot with a dense network of laterals. In monocotyledons: a developed system of adventitious roots. Roots capable of secondary thickening (in woody plants) or storage (in biennials) appear.

  4. Onset of secondary growth (in woody plants and perennial herbs). Presence of cambial stem thickening, formation of periderm (cork). This distinguishes the virginal plant from the juvenile one, whose stems are usually herbaceous, green, and lack cork.

  5. Loss of connection to seed reserves. The seedling has long since lost its cotyledons and endosperm; the plant has completely transitioned to autotrophic nutrition.

In many species, these characteristics appear not simultaneously but gradually; however, the key can be considered the moment of appearance of the first “adult” leaf after the completion of the juvenile series. In forestry practice for trees and shrubs, the beginning of the virginal stage is considered the age when the main shoot ceases to bear juvenile (e.g., lanceolate in oak) leaves and transitions to typical lobed or pinnate leaves (Gatsuk et al., 1980).

4.2. Criteria for the end of the virginal stage (transition to the generative stage)

The end of the virginal stage is marked by the transition of the plant into the generative state (g1). This transition is not always abrupt, but it can be diagnosed by the following characteristics:

  1. Appearance of generative organs (flowers, inflorescences). The most obvious sign is flowering. However, in some species (e.g., many fruit trees), flower buds are initiated a year before flowering. Therefore, a more precise criterion for the end of the virginal stage is the initiation of the first flower buds (anatomically: the transformation of the vegetative shoot apical meristem into a generative or mixed one) (Mauseth, 2016).

  2. Change in growth pattern. In many monocarpic plants (e.g., grasses), the end of the virginal stage coincides with the transition of the apical meristem from a vegetative to a generative state — the shoot stops elongating and forms an inflorescence.

  3. Reduction in vegetative growth intensity. Typically, after flowering begins, the increment of vegetative mass (leaves, stems, roots) noticeably decreases, as resources are redistributed in favor of reproductive organs.

  4. Appearance of mixed buds (in perennials). In leaf axils, along with vegetative buds, buds containing flower primordia are initiated.

  5. Change in chemical composition. The content of nitrogen (proteins) decreases in leaves and stems, while the content of carbohydrates and fiber increases.

4.3. Difficulties in differentiation: possible exceptions

In some cases, drawing a clear boundary between the virginal and generative stages can be difficult:

  • Prolonged transition (period of young generative individuals). In many tree species (oak, beech, pine), the first flowers appear on individual lower branches, while the upper part of the crown remains vegetative. Such a plant is formally generative, but functionally retains many features of the virginal stage.

  • “False” flowering. Some species, under stress conditions, may prematurely form solitary, often underdeveloped flowers, but then return to vegetative growth. This phenomenon should not be considered an exit from the virginal stage — such a transition is temporary and incomplete.

  • Vegetative reproduction during the virginal stage. In many rhizomatous and stoloniferous species, virginal individuals actively reproduce vegetatively, which externally may resemble the formation of new individuals (clones), but is not a generative process. Therefore, retention of the virginal status is not contradicted by the presence of vegetative reproduction (Gatsuk et al., 1980).

4.4. Practical use of criteria in agronomy

Knowledge of the precise criteria for the beginning and end of the virginal stage allows the agronomist to:

  • Determine the optimal harvest time for crops whose yield is formed during the virginal phase (cabbage, root crops, silage crops). Harvesting should be carried out before signs of transition to flowering appear.

  • Assess the readiness of seedlings for transplanting: virginal seedlings of fruit and vegetable crops (tomato, pepper) have higher survival rates and yield earlier harvests.

  • Diagnose “luxuriant growth” (prolongation of the virginal stage due to excess nitrogen) — absence of signs of transition to flowering despite the presence of a robust vegetative mass.

Summary of chapter:

Diagnosis of the virginal stage is based on a combination of vegetative characteristics (leaf shape, shoot and root structure) and the absence of generative organs. The onset of the stage is fixed by the completion of heteroblasty and the formation of the adult structure; the end is fixed by the initiation of the first flower buds. These criteria are universal for different life forms and serve as a reliable tool for agronomic practice.

5. Significance of the virginal stage for crop production

The virginal stage is not merely a “pause” between emergence and flowering. For the agronomist, crop scientist, forester, and breeder, this is a central period determining the quantity and quality of the harvested product. Understanding the patterns of this stage allows conscious management of plant development, shifting the balance either towards vegetative mass accumulation or towards accelerated fruiting (Yakovlev et al., 2008).

In this chapter, we will examine five key aspects of the applied significance of the virginal stage.

5.1. Formation of vegetative mass yield

For many agricultural crops, the product used by humans is formed precisely during the virginal period, before flowering and fruiting begin.

  • Forage grasses (clover, alfalfa, grasses): The maximum yield of green mass with high protein content is observed during the tillering — stem elongation phase, which corresponds to the virginal state. Mowing during this period ensures the best nutritional value of the forage. Delaying harvest leads to stem coarsening, reduced protein content, and decreased palatability (Gatsuk et al., 1980).

  • Silage crops (corn, sunflower): Harvesting for silage is carried out at the milk-wax ripeness stage of the grain, but often (especially in northern regions) corn is harvested in the virginal state — before tasseling, when the stems are maximally succulent and sugary. After transitioning to the generative phase, the stem becomes coarser, fiber content increases, and digestibility decreases.

  • Biennial vegetables (cabbage, carrot, beet): The yield of heads and root vegetables is formed exclusively during the first year of life — in the virginal stage. As soon as the plant transitions to flowering (in the second year), the storage organs become coarse, inedible, or completely disintegrate (Yakovlev et al., 2008).

  • Leafy and herb crops (lettuce, spinach, parsley, dill): Marketable product is obtained at the rosette stage, when the plant is in the virginal state. The appearance of flowering shoots (bolting) leads to loss of quality.

5.2. Managing the duration of the virginal stage: preventing “luxuriant growth”

“Luxuriant growth” is an abnormally long stay of a plant in the virginal phase without a transition to generative development (Yakovlev et al., 2008). It is often caused by excessive nitrogen nutrition combined with a deficiency of phosphorus and potassium.

  • Manifestations: Plants develop a robust, dark green vegetative mass, lodge (in cereals), but delay flowering and ripening. Seed and fruit yield sharply decreases.

  • Agronomic measures:

  • Balanced fertilization (control of nitrogen, increased proportion of phosphorus and potassium).

  • Use of growth regulators (retardants) that accelerate exit from the virginal stage and prevent lodging (e.g., chlormequat chloride on cereals).

  • Optimization of sowing dates: early sowing of winter crops ensures vernalization and timely completion of the virginal phase.

5.3. Significance for forestry and ornamental horticulture

In tree species, the virginal stage is the period of formation of the “skeleton” of the future tree (Gatsuk et al., 1980).

  • Crown formation: All major agronomic practices (trunk pruning, establishment of scaffold branches, forming a “vase” shape in fruit trees) are carried out precisely during the virginal stage. During this time, the plant is most plastic, wounds heal well, and the established shape is maintained for decades.

  • Nursery management: Saplings of fruit and forest trees are transplanted in the virginal state (2–4 years old), when they have already formed a powerful root system and a shoot typical of the species, but have not yet entered fruiting. This ensures high survival rate.

  • Forestry: The virginal stage in trees is the period of greatest competition for light and nutrients. Tending operations (thinning) in young stands are aimed at selecting the best virginal trees and forming highly productive stands.

5.4. Vegetative propagation and breeding selection

During the virginal stage, plants possess maximal regenerative capacity (Serebryakova et al., 2006).

  • Cutting propagation: Cuttings taken from virginal plants root significantly better than those from juvenile (whose tissues are not yet formed) or generative (where aging processes have already begun) plants. This is widely used in propagating currants, grapes, roses, and conifers.

  • Clonal micropropagation (in vitro): Explants (tissue fragments) from virginal plants possess a high capacity for regeneration and somatic embryogenesis (Mauseth, 2016).

  • Breeding: Selection of promising forms (hybrids) is usually carried out during the virginal stage, as vegetative characteristics (leaf shape, tillering type, habit) are already fully expressed, but there is no “interference” from flowering and fruiting.

5.5. Predicting flowering and fruiting timing

The duration of the virginal stage is a prognostic trait for assessing precocity and yield potential.

  • Precocious varieties (apple, pear, cherry) have a shortened virginal period (3–5 years compared to 10–15 in wild forms). This trait is stably inherited and used by breeders.

  • Winter crops: Passing through the virginal stage in autumn and vernalization in winter are obligatory conditions for future heading. Lack of vernalization (late sowing, warm winter) prolongs the virginal stage, leading to unproductive “luxuriant growth” and grain non-maturation.

  • Yield prediction: In field crop production, based on the condition of plants during the virginal phase (number of productive tillers, power of the root system, leaf area), future grain or fruit yield can be predicted with high accuracy.

6. Key differences from adjacent stages

For the correct diagnosis of a plant’s age state and understanding of its physiological and morphological characteristics, it is necessary to clearly distinguish the virginal stage from the preceding (juvenile) and subsequent (generative) stages. Although the transition between stages can be gradual, there are several reliable differential characteristics that allow an individual to be unambiguously assigned to a particular age group (Gatsuk et al., 1980; Serebryakova et al., 2006).

Below is a summary table of the key differences.

6.1. Comparative table of characteristics

Characteristic Juvenile stage (j) Virginal stage (v) Generative stage (g1–g3)
Capacity for flowering Absent Absent (virginal) Present (formation of flowers and fruits)
Leaf shape Juvenile (often simple, small, sometimes thread-like or lobed); heteroblasty observed Typical of the species (“adult”); heteroblasty completely ceases Typical of the species; reduced upper leaves (bracts) may appear in the inflorescence zone
Vegetative growth intensity Low or medium (depends on seed reserves) Maximal (greatest daily biomass increment) Decreases (resources redistributed to reproductive organs)
Shoot system type Simple (main shoot without branches or with weak first-order branches) Branched (skeletal axes of 2–3 orders in trees; tillering or rhizome formation in herbs) May retain complex structure, but with signs of aging (dieback of some branches)
Root system condition Primary (tap) root, lateral roots poorly developed Taproot with developed laterals formed (in dicotyledons) OR a powerful system of adventitious roots (in monocotyledons) Root system reaches maximum power, then gradually senesces (dieback of root tips)
Capacity for secondary thickening (in woody plants) Absent (stems herbaceous, green) Begins (formation of cambium, cork, lignification) Continues, but trunk growth rate slows
Response to pruning / damage Recovers weakly (limited regeneration potential) Recovers ideally (high regenerative capacity) Recovers weakly (weakened plant may die)
Nutrition type Heterotrophic (from seed) transitioning to autotrophic Fully autotrophic Fully autotrophic (with outflow of assimilates to flowers and fruits)
Capacity for vegetative reproduction Absent (or very weak) High (maximal) Reduced (especially in old generative individuals)
Examples (for tree species) Oak seedling (first 1–2 years with lanceolate leaves) Young oak (5–15 years with typical lobed leaves, but without acorns) Fruiting oak (older than 20–30 years)

6.2. Explanatory notes to the table

  • Capacity for flowering — the most reliable, but not the only characteristic. In some species, virginal individuals may form solitary sterile flowers (so-called “false flowering”), but full fruiting does not occur.

  • Growth intensity is maximal precisely during the virginal stage, as the plant does not yet expend resources on forming flowers, seeds, and fruits. This is important for practice: in forage and silage crops, harvesting should be carried out before the transition to the generative phase to obtain maximum biomass.

  • Capacity for vegetative reproduction sharply increases during the transition from the juvenile to the virginal state, which is utilized in cutting propagation and clonal micropropagation. In old generative plants, the regeneration potential decreases.

6.3. Transitional states and possible diagnostic errors

It is not always possible to draw a clear boundary between stages. Transitional forms may occur that could be mistakenly assigned to one category or another:

  1. Immature plants (im) — occupy an intermediate position between the juvenile and virginal states. They have already lost obvious juvenile characteristics but have not yet acquired all the features of an adult plant (e.g., leaves may be of an intermediate form, branching weak). They are not indicated in the table, as the article is directly devoted to the virginal stage.

  2. Young generative plants (g1) — already initiate flowers, but retain high vegetative growth intensity, which externally may resemble the virginal state. A reliable distinguishing feature is the presence of flower buds or flowers.

  3. Remontant forms (e.g., remontant raspberry, strawberry) may be simultaneously in the virginal (on young shoots) and generative (on previous year’s shoots) phases, complicating diagnosis at the whole-plant level. In such cases, assessment is carried out for each shoot individually.

6.4. Practical significance of distinguishing the stages

For the agronomist and crop scientist, the ability to distinguish age stages is necessary for:

  • Selecting the optimal harvest time (vegetative mass is harvested in the virginal stage, seeds and fruits in the generative stage).

  • Performing pruning (formative pruning — in the virginal stage, rejuvenation pruning — after productivity decline in the generative stage).

  • Collecting cuttings (best rooting from virginal plants).

  • Predicting yield (based on the condition of plants in the virginal stage, future fruiting can be assessed).

References

  1. Avila-Ospina, L., Moison, M., Yoshimoto, K., Masclaux-Daubresse, C. (2014). ‘Autophagy, plant senescence, and nutrient recycling’, Journal of Experimental Botany, 65(14), 3799-3811. doi: 10.1093/jxb/eru039 (PubMed)
  2. Evert, R.F., Eichhorn, S.E. (2013). ‘Early Development of the Plant Body’, in Raven Biology of Plants. New York: W.H. Freeman, pp. 526-537.
  3. Gatsuk, L.E., Smirnova, O.V., Vorontzova, L.I., Zaugolnova, L.B., Zhukova, L.A. (1980). ‘Age States of Plants of Various Growth Forms: A Review’, The Journal of Ecology, 68(2), 675. doi: 10.2307/2259429
  4. Lersten, N. R. (2004). ‘The Embryo’, in Flowering Plant Embryology: With Emphasis on Economic Species. Ames, Iowa: Blackwell Publishing Professional, 172-207.
  5. Mauseth, J. D. (2017). ‘Development and Morphogenesis’, in Botany: An Introduction to Plant Biology. Burlington, MA: Jones & Bartlett Learning, ch. 14.
  6. Strasburger, E., Noll, F., Schenck, H., Schimper, A. F. W. (1971). ‘Morphologie’, in von Denffer, D., Mägdefrau, K., Schumacher, W., Ehrendorfer, F. (ed.) Lehrbuch der Botanik für Hochschulen. Stuttgart: Gustav Fischer Verlag, pp. 9-202.
  7. Zotz, G., Wilhelm, K., Becker, A. (2011). ‘Heteroblasty—A Review’, The Botanical Review, 77(2), 109-151. doi: 10.1007/s12229-010-9062-8
  8. Серебрякова, Т. И., Воронин, Н. С., Еленевский, А. Г., Батыгина, Т. Б., Шорина, Н. И., Савиных, Н. П. (2006). ‘Начальные этапы онтогенеза растений Анатомия и морфология вегетативных органов [Initial stages of plant ontogenesis Anatomy and morphology of vegetative organs]’, in Ботаника с основами фитоценологии. Анатомия и морфология растений [Botany with Basic Phytocoenology. Plant Anatomy and Morphology]. Москва: ИКЦ «Академкнига», pp. 138-366.
  9. Яковлев, Г. П., Челомбитько, В. А., Дорофеев, В. И. (2008). ‘Рост, развитие и размножение [Growth, development and reproduction]’, in Ботаника [Botany]. Санкт-Петербург: СпецЛит, pp. 192-202.