Inflorescence

Last updated: June 02, 2026EspañolРусский

Inflorescence is a part of the shoot system of a plant, representing a collection of modified shoots (axes of various orders) that bear flowers and specialized leaves (bracts) and is to some degree separated from the vegetative sphere of the plant (Yakovlev et al., 2008; Serebryakova et al., 2006). Simply put, an inflorescence is a system of branching shoots designed to house and support the functions of flowers (Bell, 1991).

If in primitive flowering plants, such as magnolias or water lilies, flowers are often solitary and terminal, then during the course of evolution, most angiosperms developed groupings of flowers into inflorescences (Endress, 2010). The biological significance of the emergence of inflorescences lies in increasing the likelihood of pollination: clustering flowers makes them more visible to insect pollinators, facilitates access to nectar and pollen, and also promotes more efficient pollen dispersal in wind-pollinated plants (Yakovlev et al., 2008). Furthermore, inflorescences often ensure synchronized flower opening and protect young buds with bracts (Prenner et al., 2009). Thus, an inflorescence is not a mechanical sum of flowers, but a complex, evolutionarily established functional system that plays a key role in the plant’s reproductive success.

1. Biological and agronomic significance of inflorescences

The formation of inflorescences is not just a random clustering of flowers, but a crucial evolutionary and adaptive trait of angiosperms. The main functions of inflorescences can be divided into biological (ensuring reproductive success in nature) and agronomic (practical significance for humans).

1.1 Biological functions

The main biological function of the inflorescence is to increase pollination efficiency and, consequently, seed productivity. This function is realized through several mechanisms:

  1. Increasing visibility to pollinators. The solitary small flowers of many species (e.g., in Apiaceae or Asteraceae) would be inconspicuous to insects. When gathered into a large, conspicuous inflorescence (compound umbel or capitulum), they attract significantly more potential pollinators (Yakovlev et al., 2008). Even if individual flowers are inconspicuous, the entire inflorescence can have high attractiveness (Spitzer-Rimon et al., 2019).

  2. Facilitating access to resources. In insect-pollinated species, the inflorescence often provides a convenient “landing platform”. An insect visiting an inflorescence pollinates several flowers in a single visit, saving its time and energy, and for the plant, increases the likelihood of cross-pollination (Endress, 2010). In wind-pollinated plants (e.g., many grasses and birches), pendulous catkin inflorescences easily sway in the wind, promoting pollen dispersal (Yakovlev et al., 2008).

  3. Ensuring sequential flowering. Inflorescences often exhibit a specific order of flower opening: acropetal (from base to apex, as in a raceme) or basipetal (from apex to base, as in a dichasium). This allows the plant to extend the flowering period, guarantee cross-pollination between flowers of different ages within the inflorescence, and more efficiently allocate resources among developing fruits (Ahn, 2025; Park et al., 2012).

  4. Protection of reproductive organs. In many plants, an important protective function is performed by bracts – modified leaves in the axils of which flowers or inflorescence branches sit (Bell, 1991). They can protect young buds from desiccation, overheating, or insect herbivory. In some species (e.g., in the family Araceae), a large spathe entirely envelops the spadix inflorescence, creating a special microclimate inside (Yakovlev et al., 2008).

1.2 Agronomic significance

Inflorescences are of immense practical importance for agriculture, since in most cases they form the yield of fruits and seeds of crop plants.

  1. Yield formation. For cereals (wheat, rice, barley), vegetables (tomato, pepper, eggplant), fruits (apple, pear), and oilseeds (sunflower, rapeseed), the primary product is fruits and seeds developing from the flowers of inflorescences. The quantity and quality of these fruits directly depend on inflorescence architecture and pollination success (Sakuma et al., 2020; Ahn, 2025). For example, in bread cereals, the number of grains per spike is determined by the number and fertility of spikelets in the compound spike.

  2. Prediction and breeding. The type and structure of the inflorescence are often stable varietal characteristics, used in breeding to create high-yielding forms. For example, in tomato, inflorescence shape (simple raceme or highly branched compound inflorescence) is controlled by specific genes (such as COMPOUND INFLORESCENCE) and directly correlates with potential yield (Park et al., 2012). Assessment of inflorescence parameters (number of flowers, peduncle length, etc.) underlies physiological and breeding-based yield predictions (Dreccer et al., 2019).

  3. Practical management in agronomy. Understanding inflorescence structure is necessary for various agronomic practices:

    • Yield regulation: Pruning of tomato suckers, removal of part of the inflorescences to obtain larger fruits.

    • Selection of pollinators: For self-incompatible crops (many fruit trees: apple, pear, cherry), at least two inter-fertile varieties with coinciding flowering periods must be planted on the plot (Chabert and Mallinger, 2025).

    • Plant protection: The stages of budding and flowering are critical windows for infection by diseases (e.g., wheat loose smut enters through flowers) or damage by pests.

Thus, the inflorescence is a key organ determining not only the plant’s reproductive success in nature, but also the economic value of crops in agroecosystems.

2. Morphology of the inflorescence

To understand the vast diversity of inflorescences, one must first master their basic structure. Like any shoot, an inflorescence consists of axes, nodes, and internodes, but its structure is subordinated to the main function — optimal arrangement of flowers.

2.1 Axes of the inflorescence

The basis of any inflorescence is a system of branching axes. Two main types of axes are distinguished:

  • Main axis (axis of inflorescence, floral axis). This is the central stem from which lateral branches arise. The stem bearing the entire inflorescence is called the peduncle (Yakovlev et al., 2008; Bell, 1991). If the inflorescence consists of a single flower, its stem is usually called the pedicel (see below).

  • Lateral axes. These are branches of various orders arising from the main axis. On them, in turn, may be axes of the next order or the flowers themselves. The totality of all axes of the inflorescence forms its architecture.

Nodes and internodes on inflorescence axes are often shortened, creating a compact effect (as in a capitulum or head) or, conversely, elongated (as in a panicle or raceme) (Serebryakova et al., 2006).

2.2 Leaves of the inflorescence (bracts)

The most important difference between a reproductive shoot and a vegetative one is the presence of specialized leaves — bracts (Bell, 1991). These are modified leaves in the axils of which lateral axes or flowers develop. Bracts can differ greatly from ordinary leaves:

  • They are often smaller, may have a simple shape, be membranous, scaly, or reduced to inconspicuous outgrowths.

  • In some plants, bracts are, on the contrary, large and brightly colored, serving as attractants for pollinators (e.g., white bracts in dogwood, red bracts in poinsettia).

  • In some cases, bracts may be completely absent. Such inflorescences are called ebracteate (naked) (Serebryakova et al., 2006).

At the base of the flower there are often small leaflets — bracteoles. In dicots, there are usually two, and they are located on the sides of the pedicel (Bell, 1991).

2.3 Pedicel

The pedicel is the segment of the axis between the bract and the flower. It ends in an expansion — the receptacle — to which all parts of the flower are attached. In sessile flowers, the pedicel is absent, and the flower appears “attached” directly to the inflorescence axis (Yakovlev et al., 2008). The presence and length of pedicels determine the overall appearance of the inflorescence: in a raceme, pedicels are well-developed, while in a spike, they are barely noticeable.

2.4 Structure of the inflorescence phytomer

At the base of inflorescence morphology lies a repeating structural unit — the phytomer (sometimes called a metamer). In general terms, an inflorescence phytomer includes:

  1. A node on the axis.

  2. A bract arising from the node.

  3. An axillary structure — a lateral axis (branch of the next order) or a solitary flower.

  4. An internode up to the next node.

This repeating modular structure provides the basis for classification and understanding of inflorescence development (Park et al., 2012; Hayashi and Konishi, 1991).

2.5 Boundaries of the inflorescence

In practice, it is important to distinguish where the vegetative part of the plant ends and the inflorescence begins. Usually, the boundary is drawn at the last normally developed vegetative leaf (Bell, 1991). However, in many species there are transitional forms — from ordinary leaves at the base of the inflorescence to small bracts at its apex, which sometimes makes it difficult to precisely define the boundaries (Endress, 2010).

Thus, the inflorescence is an ordered system of axes, leaves, and flowers, where each element has its own name and function. Understanding this basic morphology is necessary for moving on to the study of inflorescence classification.

3. Classification of inflorescences (Typology)

Schematic of branching in Cannabis sativa inflorescence

Diagram of hemp inflorescence architecture (<span lang="la" class="biological-name">Cannabis sativa</span>): compound raceme and termination types in different varieties.

Shown are phytomers (internode + reduced leaf + two bracts + two solitary flowers), branching orders up to the 7th order, and terminal flower variants: in variety NB140 – terminal pistillate flower, in NB150 – hermaphroditic flower with stamens, in NB130 – indeterminate inflorescence without terminal flower.

The diversity of inflorescences follows certain patterns of branching. The modern classification is based on two main characteristics: the mode of axis growth (monopodial or sympodial) and the degree of branching (simple or compound). Accordingly, two main types are distinguished: botryoid (racemose) and cymoid (cymose) (Endress, 2010; Yakovlev et al., 2008).

3.1 Botryoid (racemose) inflorescences

This type is characterized by monopodial growth of the main axis: the apical bud functions throughout the growth period of the inflorescence and does not terminate in a flower (except in rare cases – see below). Flowers or lateral inflorescences are initiated on the main axis in acropetal order (from base to apex), meaning the oldest flowers are at the base, and the youngest at the apex (Endress, 2010; Serebryakova et al., 2006).

Botryoid inflorescences are divided into simple (flowers are borne directly on the main axis) and compound (the main axis bears not individual flowers, but partial inflorescences).

Simple botryoid inflorescences:

Spike of orchid Ophrys insectifera

Spike inflorescence of the orchid <span lang="la" class="biological-name">Ophrys insectifera</span>. Sessile flowers are sequentially arranged on an elongated main axis.

In a spike, unlike a raceme, pedicels are absent (sessile flowers). This is one of the most common inflorescence types in wind-pollinated plants, as well as in some insect-pollinated ones (as in this example).

Corymb of yarrow

Corymb inflorescence of yarrow <span lang="la" class="biological-name">Achillea filipendulina</span>. Flowers are positioned almost in one plane due to different pedicel lengths.

In a corymb, lower pedicels are longer than upper ones, creating a “flat-topped” appearance. This structure is characteristic of many members of the Rosaceae family, e.g., pear and rowan.

  • Raceme. Main axis elongated, flowers on well-developed pedicels of approximately equal length. Examples: bird cherry, lupine, cabbage (in seed).

  • Corymb. A variant of a raceme with a shortened main axis and unequal pedicel length: lower pedicels longer than upper ones, so that all flowers are at approximately the same level. Example: pear, spirea (Yakovlev et al., 2008).

  • Spike. Main axis elongated, but flowers sessile (without pedicels). Example: plantain, orchis. In grasses, the compound spike (see below) is the most important inflorescence of cereal crops (Sakuma et al., 2020).

  • Spadix. A spike with a greatly thickened, fleshy main axis. Often surrounded by a large bract – a spathe. Characteristic of aroids (calla lily), as well as maize, where the female inflorescences are cobs (Bell, 1991).

  • Catkin. A pendulous spike (rarely a raceme), usually bearing unisexual flowers and falling as a whole after fruiting. Typical of wind-pollinated trees (birch, willow, poplar) (Yakovlev et al., 2008).

  • Head. Main axis greatly shortened and expanded, flowers sessile or on very short pedicels, gathered in a dense cluster. Example: clover, alfalfa.

  • Capitulum. A specialized head with a flattened, dish-shaped receptacle on which sessile flowers are densely arranged. The capitulum is surrounded externally by an involucre of modified bracts. This is the characteristic inflorescence of the family Asteraceae (Compositae) – sunflower, aster, dandelion (Ahn, 2025; Yakovlev et al., 2008).

  • Umbel. Pedicels of approximately equal length arise from the apex of a shortened main axis. A simple umbel is characteristic of cherry, primrose.

Compound botryoid inflorescences:

Panicle diagram

Diagram of a panicle — a compound branching inflorescence with a monopodial main axis and branching lateral axes.

The illustration shows the main axis, from which lateral branches of different orders arise, bearing flowers on pedicels. Branching degree decreases from base to apex, creating a characteristic pyramidal shape.

In these inflorescences, not flowers but partial inflorescences (which have the structure described above) develop on the main axis.

Compound umbel of Lomatium utriculatum

Compound umbel inflorescence of <span lang="la" class="biological-name">Lomatium utriculatum</span>. The rays of the compound umbel and the umbellets at their ends are clearly visible.

In a compound umbel, rays arise from the apex of the main axis, and each ray bears a simple umbel (umbellet) at its tip. Such architecture is characteristic of many members of the Apiaceae family (Umbelliferae, Celery family).

  • Panicle. A compound, repeatedly branched inflorescence in which the degree of branching of lateral axes decreases from base to apex (pyramidal form). Partial inflorescences are simple racemes or spikes. Examples: lilac, grape, oat, rice (Dreccer et al., 2019; Park et al., 2012). In grasses, oats, millet, and rice have a panicle.

  • Compound spike. The main axis bears sessile partial inflorescences – spikelets, each of which is a simple spike (in grasses, of several flowers covered by scales). This is the typical inflorescence of wheat, rye, barley (Sakuma et al., 2020).

  • Compound umbel. Partial inflorescences (simple umbels called umbellets) arise from the apex of the main axis on rays of unequal length. Characteristic of the Apiaceae family (Umbelliferae) – dill, carrot, coriander.

  • Compound corymb. Partial inflorescences (short racemes or corymbs) are arranged on the main axis such that they all lie in approximately the same plane. Example: rowan, viburnum.

3.2 Cymoid (cymose) inflorescences

Dichasium diagram

Diagram of a dichasium — a cymose inflorescence with two opposite lateral branches below the terminal flower.

The principle of sympodial growth is shown: the terminal flower opens first, then the flowers on the lateral axes. Such a structure is characteristic of many members of the Caryophyllaceae family.

This type is characterized by sympodial growth: the main axis ends in a flower, and further growth of the inflorescence occurs through one or more lateral axes, which behave similarly. Flowers open in basipetal order (from top to bottom), i.e., the terminal flower of the main axis opens first (Endress, 2010; Serebryakova et al., 2006).

Cymoid inflorescences are usually parts of more complex structures (e.g., thyrses) and are rarely found in pure form as the entire inflorescence. Main types:

  • Dichasium. Under the terminal flower, two opposite lateral axes develop, which also end in flowers and can branch similarly. The result is a forked inflorescence. Example: chickweed, carnation.

  • Monochasium. After flowering of the main axis, only one lateral axis develops, which also ends in a flower. Depending on the arrangement of subsequent flowers, helicoid cyme (flowers arranged spirally) and scorpioid cyme (flowers arranged zigzag in one plane) are distinguished. Examples: forget-me-not, gladiolus (Bell, 1991; Hayashi and Konishi, 1991).

3.3 Mixed and aggregate inflorescences

Diagrams of cymose inflorescences

Classical diagrams of cymose inflorescences from the botanical atlas by Gustav Hegi (1906).

The illustrations show various types of monochasia and dichasia, as well as their combinations. Such diagrams help understand the principles of sympodial branching and the sequence of flower opening.

In nature, inflorescences combining traits of both botryoid and cymoid types are often found. The most well-known is the thyrse – a main axis that grows monopodially (as in a raceme), but the lateral inflorescences on it are cymose (dichasia or monochasia) (Endress, 2010). The thyrse is widespread, for example, in Lamiaceae (mint, sage), Solanaceae (tomato, potato), Scrophulariaceae. Complex inflorescences built from elementary capitula, umbels, or spikelets are often called aggregate: “panicle of capitula” (in some Asteraceae), “panicle of umbels” (in Araliaceae) (Yakovlev et al., 2008).

3.4 Principles of development and evolution

From an ontogenetic point of view, botryoid inflorescences with indeterminate growth are considered more primitive. Cymoid inflorescences generally arise secondarily as a result of specialization to particular pollination types. It has been established that within a single inflorescence, a transition from botryoid branching at the base to cymoid at the apex (so-called thyrses) is evolutionarily more advanced (Endress, 2010). Understanding these patterns is necessary not only for taxonomists but also for breeders, since inflorescence architecture in many crops (tomato, sunflower, cereals) is directly linked to productivity (Park et al., 2012; Ahn, 2025).

4. Anatomical structure of the inflorescence

The inflorescence, like any other plant organ, consists of various types of tissues that provide its strength, transport of nutrients, and protection. Although the general structural plan of inflorescence axes is similar to that of the vegetative stem, it has a number of features related to the performance of the reproductive function.

4.1 Conducting system

The axes of the inflorescence must provide intensive transport of water, mineral salts, and, mainly, organic substances (assimilates) to developing flowers and fruits. The conducting system is represented by vascular bundles that run along all axes.

  • Xylem (from Greek xylon – wood) provides the upward flow of water and minerals from the roots to the flowers and leaves of the inflorescence.

  • Phloem (from Greek phloos – bark, bast) provides the downward flow of organic substances, mainly sucrose, from photosynthetic leaves to growing flowers and ovules (Serebryakova et al., 2006).

An important feature of inflorescences is the strong development of vascular bundles in the zones of attachment of flowers and branches, especially at the nodes. This ensures efficient supply of each element of the inflorescence. In many species, vascular “knots” form at the sites of pedicel attachment, which can serve as zones of nutrient retention, influencing the order of flower opening and fruit development (Park et al., 2012). In crops with large inflorescences and fruits (sunflower, tomato, maize), the conducting system of the peduncle is particularly well developed to deliver large amounts of assimilates to the developing yield (Ahn, 2025).

4.2 Mechanical tissues

Inflorescences, especially large and heavy ones (e.g., sunflower, maize cobs, millet panicle), must support their own weight and the impact of wind and rain. This function is performed by mechanical tissues, primarily collenchyma and sclerenchyma.

  • Collenchyma is located directly under the epidermis and consists of living cells with unevenly thickened walls. It provides strength to young, growing inflorescence axes.

  • Sclerenchyma consists of dead cells with evenly thickened, often lignified walls (sclereids or fibers). It forms a continuous cylinder or strands in the cortex and pith, providing exceptional resistance to bending and compression (Yakovlev et al., 2008).

Sclerenchyma is especially well developed in the axes of grass inflorescences (culm), where it forms a ring of mechanical tissue, as well as in the pedicels and receptacles of Asteraceae capitula. In wind-pollinated species with long, thin catkins, mechanical tissues ensure their flexibility and resistance to wind (Endress, 2010).

4.3 Protective tissues and secretory structures

The surface of inflorescence axes and bracts is covered by epidermis. The epidermis often bears a cuticle – a waxy layer that protects against desiccation and wetting. Also on the surface may be trichomes (hairs) and glands.

  • Trichomes can perform a protective function (against insect herbivory, overheating) and participate in creating a microclimate around young buds.

  • Glandular hairs and nectaries are often located on the inflorescence axes and bracts. They secrete nectar, which attracts insect pollinators. In some plants (e.g., cotton), there are extrafloral nectaries on the bracts and pedicels (Yakovlev et al., 2008; Spitzer-Rimon et al., 2019).

In many species, the tissues of the inflorescence contain idioblasts – specialized cells containing tannins, essential oils, alkaloids, or calcium oxalate crystals. These substances can repel herbivores and protect the inflorescence from pathogens (Bell, 1991).

4.4 Formation of abscission zones

An important anatomical feature of pedicels and fruit stalks is the presence of an abscission (separation) layer at their base. This consists of several rows of small, thin-walled, often lignified cells that break or soften when fruits ripen or flowers wither, leading to the detachment of the fruit or faded flower (Ahn, 2025). In cereal crops, the strength of grain retention in the spike (resistance to shattering) is determined precisely by the structure of the abscission layer at the base of the spikelets.

Conclusion

The anatomical structure of the inflorescence is a complex of interconnected tissues specialized to perform three main tasks: conducting substances, providing mechanical strength, and protecting the reproductive organs. Features of inflorescence anatomy (vascular bundles, mechanical fibers, abscission zones) are of great practical importance, as they determine lodging resistance, ease of threshing, and transportability of the crop.

5. Formation and development (Ontogenesis of the inflorescence)

Wheat inflorescence development

Stages of wheat inflorescence development: from double ridge to white anther stage.

The diagram shows: (a) double ridge stage – formation of spikelet and leaf primordia; (b) terminal spikelet stage; (c) white anther stage – active function of spikelet meristem and flower formation.

The development of the inflorescence is a regular process of transformation of vegetative growing points into generative ones, which includes several sequential stages. Understanding the ontogenesis of the inflorescence is necessary for controlling flowering time, predicting yield, and interpreting phenological observations in crop production.

5.1 Transition from vegetative to reproductive growth

Initiation of the inflorescence begins with the transformation of the vegetative shoot apical meristem into an inflorescence meristem (IM). This transition is controlled by a complex of endogenous (plant age, hormonal status) and exogenous factors (photoperiod, temperature, nutrition) (González-Suárez et al., 2025).

In most plants, the first visible sign of transition is an increase in size and a change in shape of the apical meristem – it becomes wider and more convex (“umbrella” or “dome” shape). In grasses, this transition is morphologically expressed in the “double ridge” stage – two types of primordia appear on the apical meristem: the primordium of the future spikelet and an underdeveloped leaf primordium (Sakuma et al., 2020; Dreccer et al., 2019). In dicots, such as tomato, the transition from vegetative to generative meristem is also accompanied by its enlargement and a change in the pattern of leaf primordium initiation (Park et al., 2012).

A key role in this switch is played by hormonal and genetic factors, particularly the interaction of florigen (FT protein) and anti-florigenic systems (e.g., TERMINAL FLOWER1 in Arabidopsis), which integrate environmental signals (Landrein et al., 2025). In winter and biennial crops, a necessary condition for the transition is vernalization – exposure to low temperatures, which suppresses the expression of flowering repressor genes (e.g., FLC in Brassicaceae) (González-Suárez et al., 2025).

5.2 Sequence of inflorescence element initiation

After transformation into an inflorescence, the apical meristem begins to form axes and flowers in a specific sequence. Two main types of initiation are distinguished (Serebryakova et al., 2006):

  1. Acropetal (from Greek akron – tip, petra – stone). Flowers or lateral inflorescences are initiated first at the base of the inflorescence, and the youngest at the apex. This type is characteristic of botryoid (racemose) inflorescences – raceme, spike, panicle. Flower opening during acropetal development usually occurs from base to apex (centripetally with respect to the entire inflorescence).

  2. Basipetal (from Greek basis – base, petra – stone). The terminal flower of the main axis is initiated first, followed sequentially by lateral flowers or axes from top to bottom. This type is characteristic of cymoid (cymose) inflorescences – dichasia, monochasia. Flower opening proceeds from top to bottom.

In complex inflorescences, such as the thyrse, the acropetal order may be maintained on the main axis (growing monopodially), while the lateral cymose branches have a basipetal order (Endress, 2010). Understanding the type of initiation helps the agronomist assess the degree of inflorescence development at different stages.

5.3 Phenological phases of inflorescence development in major crops

In agronomy, systems of phenological scales (e.g., the Zadoks scale for cereals, the BBCH scale for a wide range of crops) are used to detail the stages of inflorescence formation. Key phases:

  1. For cereal crops (wheat, barley, rye, oat, rice):

    • Stem elongation. Beginning of rapid internode growth, elevation of the apical meristem above the soil surface. Spike differentiation occurs inside the stem.

    • Heading. Emergence of the upper part of the inflorescence (spike or panicle) from the leaf sheath (Dreccer et al., 2019). This phase is critically important as it determines susceptibility to diseases (e.g., loose smut infects precisely during heading).

    • Flowering. Anther extrusion, flower opening, and pollination. In wheat, flowering often begins with flowers in the middle part of the spike (Sakuma et al., 2020).

  2. For legumes and oilseed crops (pea, soybean, rapeseed, sunflower):

    • Bud formation. Appearance of visible inflorescence buds (raceme, capitulum).

    • Beginning of flowering. Opening of the first flowers in the inflorescence. In complex inflorescences (sunflower capitulum), flowering begins with the ray florets and proceeds toward the center (centripetally) (Ahn, 2025).

  3. For fruit and vegetable crops (tomato, apple, grape):

    • Inflorescence differentiation. Formation of inflorescence primordia in generative buds (usually in the previous year).

    • Inflorescence emergence (in grape). Growth and branching of the inflorescence (panicle) after bud break.

5.4 Molecular-genetic basis of inflorescence development (concept of “maturation clock”)

Modern research, especially in tomato, has shown that inflorescence development is governed by a “meristem maturation clock” (Park et al., 2012). During development, the apical meristem passes through a series of sequential states, each corresponding to a specific set of transcription factors. In tomato and other solanaceous plants that grow sympodially, the inflorescence forms as a result of the meristem, after passing through a stage of forming several leaves, terminating in a flower. The COMPOUND INFLORESCENCE (S) gene and other regulators (FALSIFLORA, ANANTHA) control the rate of this maturation (Park et al., 2012). Mutations that slow down maturation lead to the formation of more branched inflorescences with a greater number of flowers.

In Arabidopsis and related species, genes of the MADS-box family (SOC1, SVP, APETALA1) and the LEAFY protein play crucial roles in switching from vegetative growth to flower formation (González-Suárez et al., 2025). The WUSCHEL–CLAVATA regulatory circuit controls the maintenance of the stem cell population and determines meristem size, which in turn affects the number of flowers in the inflorescence (Landrein et al., 2025). In cereal crops, genes similar to FRIZZY PANICLE (FZP) control the transition from formation of inflorescence branches to formation of spikelets, determining the final architecture of the compound spike or panicle (Sakuma et al., 2020).

Thus, inflorescence formation is a strictly programmed process involving a switch in meristem fate, ordered initiation of elements, and is governed by a complex network of regulatory genes. Knowledge of inflorescence ontogenesis and its genetic basis allows breeders and agronomists to purposefully influence yield architecture and adapt varieties to specific conditions.

6. Influence of environmental factors on the inflorescence (Ecological plasticity)

Inflorescence architecture and its reproductive success are not rigidly determined traits. They are shaped by a complex of abiotic factors (light, temperature, humidity, mineral nutrition), which can cause significant morphological changes — from variation in flower number to complete structural transformation. This ability of the inflorescence to alter its parameters in response to environmental conditions is called ecological plasticity.

6.1 Light and photoperiod

Light is a key signaling factor controlling the transition to flowering and subsequent inflorescence development.

  • Photoperiodic sensitivity. In many species (especially plants of temperate latitudes), inflorescence initiation is induced by a specific day length. For example, in the short-day plant Cannabis sativa, under long-day conditions (16 h) solitary flowers form in leaf axils, but upon transition to short days (12 h) intense branching occurs, forming compound racemose inflorescences (Spitzer-Rimon et al., 2019). Thus, photoperiod influences not only flower initiation but also inflorescence branching architecture.

  • Light intensity and quality. Low light or shading generally leads to elongation of internodes in the inflorescence (etiolation), which can reduce its compactness. High far-red (FR) light content, characteristic of shaded conditions, in many species stimulates inflorescence branching and can cause “vegetative reversion” — transformation of flowers into vegetative shoots (Dreccer et al., 2019; González-Suárez et al., 2025).

6.2 Temperature

Temperature conditions during inflorescence formation affect the rate of development, number of flowers, and their fertility.

  • Effect on phenology. Elevated temperature accelerates inflorescence development and maturation, shortening the time from budding to flowering. This can lead to a reduction in flower number due to faster progression through initiation phases (Dreccer et al., 2019). Low temperatures, conversely, slow growth and may promote an increase in flower number per inflorescence, as the meristem remains in the vegetative phase longer (González-Suárez et al., 2025).

  • Temperature stresses. High temperatures (heat) during generative organ formation often lead to pollen sterility, manifesting as reduction of apical flowers (abortion) and reduced seed set per fruit (Chabert and Mallinger, 2025; Sakuma et al., 2020). In cereals, heat stress during meiosis causes spikelet sterility. Low freezing temperatures (frosts) during flowering can damage flowers, leading to empty grains and crop failure.

  • Vernalization. In winter and biennial crops, a prolonged exposure to low temperatures (vernalization) is required for the transition to inflorescence formation. Without such treatment, the plant remains in a vegetative state (González-Suárez et al., 2025).

6.3 Mineral nutrition

The supply of mineral nutrients, primarily nitrogen and phosphorus, strongly influences inflorescence parameters.

  • Nitrogen. High soil nitrogen content stimulates vegetative growth but can also increase inflorescence size (raceme length, flower number) in many crops, if not exceeding a certain threshold (Landrein et al., 2025). However, excess nitrogen often leads to excessive internode elongation, stem lodging, and reduced inflorescence strength. Nitrogen deficiency, on the contrary, reduces flower number and causes premature cessation of inflorescence development.

  • Phosphorus. Phosphorus deficiency is critical at the stage of flower initiation. Phosphorus starvation slows inflorescence formation, reduces flower number, and lowers fertility (number of set seeds) (Chabert and Mallinger, 2025).

  • Effect on branching. In tomato and other solanaceous plants, improved mineral nutrition can enhance inflorescence branching by extending the period of meristem activity (Park et al., 2012). Conversely, nutrient deficiency often leads to reduction of lateral inflorescence branches.

6.4 Water regime and drought

Water stress is one of the strongest factors limiting reproductive development.

  • Flower reduction. Drought during budding and flowering causes massive flower abortion, especially in the upper (apical) part of the inflorescence, where the youngest, most competitive flowers are located (Hayashi and Konishi, 1991). This phenomenon is well studied in cereals (wheat, maize) and legumes (soybean).

  • Ovary abortion. With prolonged water deficit, the processes of pollination and fertilization can be disrupted, and developing ovaries may die. This manifests as the formation of “empty” fruits without seeds (Chabert and Mallinger, 2025).

  • Structural changes. In some species, under moisture deficiency, inflorescences become more compact, with shortened internodes, which is an adaptive response to reduce transpiration (Doidy et al., 2024).

6.5 Phenomenon of vegetativization (prolification) of the inflorescence

Under the influence of unfavorable factors (long day, elevated temperatures, injury) or mutations, vegetativization (or prolification) of the inflorescence may occur — a reverse development in which flowers turn into vegetative shoots (leaves, stems). In this case, the inflorescence axis continues to grow, and rosettes of leaves or new vegetative shoots form instead of flowers (González-Suárez et al., 2025; Endress, 2010). This phenomenon is often observed in roses, brassicas (cauliflower, broccoli), and some grasses, and can be used in breeding (e.g., creating giant inflorescences in cauliflower).

7. Practical management and agronomic significance

Knowledge of inflorescence morphology, ontogenesis, and ecological plasticity underlies many agronomic and breeding practices aimed at increasing yield, improving product quality, and enhancing plant stress tolerance.

7.1 Regulation of load and inflorescence architecture

A key agronomic practice is regulating the number of flowers and ovaries in the inflorescence to optimize fruit size and quality.

  • Pruning (removal of lateral shoots). In tomato, pepper, eggplant, and other solanaceous crops, removal of excess vegetative shoots (suckers) redirects assimilates to the remaining inflorescences. A more drastic operation — removal of part of the inflorescences (crop load regulation) — allows obtaining larger and more uniform fruits (e.g., in large-fruited tomato, pepper varieties) (Park et al., 2012).

  • Topping (removal of inflorescence apex). In maize, removal of the male inflorescence (tassel) at the tasseling stage (topping) stimulates assimilate flow to the developing ears and is used to produce seed maize and prevent unwanted cross-pollination. In ornamental crops, topping the main shoot induces inflorescence branching (e.g., in chrysanthemum).

  • Use of growth regulators. Retardants (e.g., chlormequat chloride, paclobutrazol) inhibit cell elongation, leading to shortening of inflorescence internodes, increased stem strength (lodging resistance), and sometimes increased flower number (Landrein et al., 2025). Flowering stimulants (gibberellins) can induce inflorescence development in some crops under unfavorable conditions, but their use must be cautious, as they can cause vegetativization (prolification) (González-Suárez et al., 2025).

7.2 Yield prediction and breeding

Inflorescence architecture serves as a reliable morphological marker for selecting high-yielding genotypes.

  • Assessment of yield components. In cereal crops, direct counting of the number of productive spikes per unit area, number of spikelets per spike, and number of grains per spikelet underlies yield structure calculations (Sakuma et al., 2020). In sunflower, the number of achenes per capitulum correlates with the capitulum diameter, which is easy to measure. In tomato and legumes, the number of flowers per inflorescence (raceme) is a predictive trait of potential yield (Park et al., 2012).

  • Breeding for desired architecture. Genes controlling inflorescence branching are targets for breeding. For example, in tomato, alleles of the COMPOUND INFLORESCENCE (S) gene, which cause formation of compound branched inflorescences, have been fixed in some varieties to increase yield (Park et al., 2012). In cereals, genes responsible for spikelet number (FRIZZY PANICLE, etc.) are used to create forms with more productive compound spikes or panicles (Sakuma et al., 2020; Ahn, 2025).

  • Use of heterosis. For many cross-pollinated crops (rye, sunflower, maize), the production of hybrid seeds requires an effective system for controlling pollination. Understanding the structure of male and female inflorescences, as well as the phenology of their flowering, is critically important for organizing hybridization (Chabert and Mallinger, 2025).

7.3 Pollination management and selection of pollinator varieties

For crops with partial or complete self-incompatibility (apple, pear, cherry, many Brassicaceae, some tomato varieties), cross-pollination must be ensured.

  • Selection of pollinator varieties. When establishing an orchard or sowing a field, it is important to place inter-fertile varieties in close proximity to each other. The flowering periods of the varieties must coincide, and their inflorescences must be compatible with respect to genetic self-incompatibility (SI) or early inbreeding depression (EID) factors (Chabert and Mallinger, 2025). For example, for the apple variety ‘Antonovka’, suitable pollinators with simultaneous flowering are ‘Anis’, ‘Shtreifling’.

  • Use of pollinators. In protected cultivation (greenhouses), bumblebees (Bombus terrestris) or honeybees are used for pollination of tomatoes, peppers, and eggplants. Pollination efficiency directly depends on inflorescence architecture: in tomato, the raceme should be open and accessible to insects (Dreccer et al., 2019).

  • Phenological forecasts. Knowledge of inflorescence development phases (budding, flowering) allows timely application of fertilizers, irrigation, pest control treatments, and accurate determination of harvest dates. For stone fruit crops (cherry, apricot), it is important to forecast flowering dates to choose optimal timing for protective measures against frost.

7.4 Protection of inflorescences from diseases and pests

Inflorescences are the most vulnerable plant organs, as they serve as “entry points” for many pathogens.

  • Fungal diseases. Loose smut of cereals (Ustilago tritici, U. nuda) infects the inflorescence at the heading stage, turning the spike into a mass of dark spores. Septoria and Fusarium head blight cause death of flowers and grains. Fungicide treatments must be carried out strictly at the stem elongation and early heading stages (Sakuma et al., 2020).

  • Bacterial and viral diseases. Fire blight of fruit crops penetrates through flowers, causing wilting and blackening of inflorescences. Tobacco mosaic virus (TMV) infects tomatoes through pedicels.

  • Pests. Apple blossom weevil (Anthonomus pomorum) lays eggs in buds, causing them to dry out. Thrips and aphids often concentrate in inflorescences. Agronomic monitoring of the budding phase is necessary for timely application of insecticides.

  • Use of resistance. Breeding varieties with dense, closed inflorescences (e.g., cabbage – head) that are less accessible to pests and pathogens is an important direction.

7.5 Stress impacts and adaptation measures

Knowledge of inflorescence ecological plasticity allows the development of strategies to reduce stress damage.

  • Adaptation to drought. In drought-tolerant wheat and barley varieties, the inflorescence (spike) often has a waxy bloom, small scales, and the ability to abort the least valuable (apical) flowers under water deficit (Hayashi and Konishi, 1991; Sakuma et al., 2020).

  • Lodging resistance. Short, compact inflorescences with thick peduncles and well-developed mechanical tissue (sclerenchyma) are less prone to lodging. This trait is selected for in cereals and ornamental crops (Ahn, 2025).

  • Use of stress protectants. Treatment of plants with salicylic acid, brassinosteroids, or abscisic acid at the budding stage can increase inflorescence thermo- and drought tolerance, reducing flower abortion (Chabert and Mallinger, 2025; Landrein et al., 2025).

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