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Morphological and anatomical adaptations to water regime

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Morphological and anatomical adaptations of plants to water regime are a set of hereditarily fixed external (morphological) and internal (anatomical) features of vegetative organs that ensure the maintenance of water balance under conditions of varying environmental moisture supply (soil and atmospheric).

Unlike plant physiology, which studies the movement of water and solutes at the cellular and tissue levels, ecological morphology focuses on macro- and microscopic structures that have been formed in the course of evolution as a response to constant or seasonal action of a particular factor. These structures allow the plant either to limit water loss (when water is deficient) or, conversely, to actively remove water and ensure gas exchange when it is in excess (Strasburger et al., 1971; Serebryakova et al., 2006).

The ability of plants to adapt to water regime is expressed in the formation of two opposite directions of organ structure:

  • Xeromorphosis (xeromorphic structure) — a complex of traits that arise under conditions of water deficit (drought, dry air, soil freezing) and aim to reduce transpiration, store water, or enhance its uptake (Strasburger et al., 1971; Beck, 2010). Typical examples: small, stiff leaves, thick cuticle, sunken stomata, dense pubescence, development of water-storage parenchyma (succulents).

  • Hydromorphosis (hydromorphic structure) — a complex of traits that arise under conditions of constant water excess (aquatic environment, waterlogged soils) and aim to facilitate gas exchange and water removal (Serebryakova et al., 2006; Roland & Roland, 1980). Typical examples: thin and soft leaves with large intercellular spaces (aerenchyma), poorly developed mechanical tissue, reduction of cuticle and stomata.

In relation to water regime, all plants are conventionally divided into three main ecological groups, each demonstrating its own spectrum of morphological and anatomical adaptations (Yakovlev et al., 2008; Serebryakova et al., 2006):

  1. Hygrophytes — plants of excessively moist habitats;

  2. Mesophytes — plants of moderate moisture (they occupy an intermediate position; most cultivated plants belong to this group);

  3. Xerophytes — plants of arid habitats (including succulents and sclerophytes).

It is important to understand that adaptations to water regime rarely occur in isolation. They are closely linked to adaptations to light and temperature regimes, which leads to the formation of the plant’s holistic appearance — its life form (Raunkiaer, 1934; Niklas, 2008). The following sections of the article will be devoted to a detailed examination of the specific morphological and anatomical features of each of the identified ecological groups.

1. Theoretical basis: target organs of adaptations

Wax cuticle on a geranium leaf
©Rosser1954commons.wikimedia.orgCC BY-SA 4.0

Wax cuticle on the surface of a geranium leaf

Macrophotograph clearly showing the waxy layer (cuticle) that acts as a barrier against water evaporation.

Before considering specific ecological groups, it is necessary to understand which organs and tissues of plants are the main “targets” for adaptive changes in response to altered water regime. Since water is involved in maintaining cell shape (turgor), serves as a medium for metabolic reactions, and is an electron donor for photosynthesis, its deficit or excess primarily affects the functioning of those structures responsible for gas exchange, photosynthesis, transport and mechanical strength (Strasburger et al., 1971; Evert, 2006).

A comparative analysis of the morphology and anatomy of plants from different ecological groups shows that all adaptive changes affect the same four categories of organs and tissues, which can be called target organs.

1.1. The leaf as the main “water cost” centre

The leaf is the main transpiration organ. Up to 95% of the water absorbed by the plant evaporates through its surface (especially through stomata). Therefore, the anatomical structure of the leaf blade is the most informative indicator of environmental conditions (Beck, 2010; Evert, 2006).

  • Epidermis: In dry conditions, epidermal cells acquire thickened outer walls, are covered with a thick cuticle and waxy layer (the cuticle may be folded, which increases the evaporation area during sharp fluctuations in humidity). In humid conditions, the cuticle is thin or absent, and cell walls are thin (Roland & Roland, 1980).

  • Stomatal apparatus: The number, size, depth of stomata and the presence of subsidiary cells vary. In xerophytes, stomata are often sunken in depressions (crypts), covered with hairs or wax plugs, which reduces the evaporation rate. In hydrophytes, stomata are located on the upper side of floating leaves or are completely absent (Evert, 2006; Serebryakova et al., 2006).

  • Mesophyll (photosynthetic parenchyma): Under conditions of good lighting and water deficit, palisade (columnar) parenchyma forms in one or several layers, which increases the efficiency of photosynthesis under limited transpiration. Under excess moisture and shade, loose spongy parenchyma with large intercellular spaces that provide gas exchange predominates (Yakovlev et al., 2008).

  • Conducting and mechanical tissues of the leaf: In the veins of xerophytes, xylem (water-conducting vessels) and sclerenchyma (supporting fibres) are more strongly developed. In hydrophytes, conducting bundles are reduced, and mechanical tissues are weakly developed or absent (Strasburger et al., 1971).

1.2. The stem: support and reserve channel

The stem performs supporting and conducting functions. In arid conditions, the proportion of sclerenchyma and lignified xylem elements increases, which gives the stem rigidity and protects it from lodging when turgor is lost (so-called “sclerophyllization”). In many xerophytes, the function of photosynthesis is taken over by green stems (cladodes, phylloclades), and leaves are reduced (Beck, 2010). In humid conditions and aquatic plants, stems contain powerful aerenchyma — parenchyma with large air-bearing cavities, which provides buoyancy and oxygen supply to submerged parts (Roland & Roland, 1980).

1.3. The root system: water uptake and retention

The root is the organ that absorbs water and mineral salts. Although its structure depends less on air humidity, it reflects the water regime of the soil. In xerophytes, root systems can be exceptionally powerful (taproots reaching great depths) or, conversely, superficial but highly branched (to collect condensate and rainwater). In the root anatomy of xerophytes, xylem and pericycle are well developed, and an additional suberized exodermis is often present, reducing water loss from the cortex. In aquatic plants, the root system is reduced, root hairs are almost absent, and roots may be replaced by rhizoids or completely absent (Yakovlev et al., 2008).

1.4. Special tissues and structures

  • Aerenchyma: Specialized air-bearing parenchyma with large intercellular spaces, characteristic of hydrophytes and hygrophytes. It facilitates gas exchange under conditions of oxygen deficiency (underwater, in waterlogged soil) (Serebryakova et al., 2006).

  • Hydathodes: Water stomata or special glands that secrete liquid water droplets (guttation). Characteristic of hygrophytes living in high air humidity conditions when transpiration is hindered (Evert, 2006).

  • Trichomes (hairs): Perform many functions: reflection of sunlight (reducing heating), creation of a boundary layer of still air (reducing transpiration), protection from insects, and in some desert plants, absorption of water from dew or fog (Beck, 2010).

All these structures change not in isolation but as a single functional system. In the next section, we will examine how these changes specifically manifest in hygrophytes, mesophytes and xerophytes.

2. Group 1: Hydrophytes (Aquatic plants)

Hydrophytes (from Greek hydor — water and phyton — plant) are an ecological group of higher plants that live in an aquatic environment or under conditions of constant excessive soil moisture, with their vegetative bodies fully or partially submerged in water. Strictly speaking, the term “hydrophytes” refers to plants that are rooted at the bottom and raise part of their shoots above water, while fully submerged forms are called hydatophytes, and free-floating ones — pleistophytes (Scremin-Dias, 2009; Yakovlev et al., 2008). However, in educational literature, they are often combined into one ecological group due to the similarity of their main adaptations.

For hydrophytes, water is not only a habitat but also a limiting factor, but with the opposite sign: instead of water deficit, they face its excess, which creates two main problems:

  1. Difficult gas exchange: the solubility of oxygen and carbon dioxide in water is significantly (tens of times) lower than in air. As temperature rises, gas solubility drops even more, which can lead to tissue hypoxia (Iftikhar et al., 2025).

  2. Lack of mechanical support: water supports the plant, removing the need for powerful mechanical tissues (collenchyma and sclerenchyma), but at the same time creates resistance during currents and waves.

Consequently, the anatomical and morphological adaptations of hydrophytes are aimed at maximally increasing the gas exchange surface, facilitating oxygen transport to submerged organs, and reducing the cost of building supporting structures.

2.1. Classification of hydrophytes by life forms

With respect to the aquatic environment and substrate, several life forms are distinguished among hydrophytes (Scremin-Dias, 2009; Serebryakova et al., 2006):

  • Submerged plants: completely underwater (e.g., Elodea canadensis, Ceratophyllum demersum, Myriophyllum spicatum). They may lack stomata, and absorption of water and mineral salts occurs over the entire body surface.

  • Plants with floating leaves: rooted at the bottom, but their leaf blades are held on the water surface (e.g., Nuphar lutea, Nymphaea alba, Victoria amazonica). Floating leaves exhibit heterophylly — differences in the structure of submerged and emergent leaves.

  • Free-floating plants: not attached to the bottom, drift in the water column or on its surface (e.g., Lemna minor, Salvinia natans, Eichhornia crassipes). They often show reduction of roots and predominance of vegetative reproduction.

  • Amphibious plants: able to grow both in water and in air, changing leaf anatomy depending on the hydroperiod phase (e.g., Polygonum amphibium, Sagittaria sagittifolia) (Scremin-Dias, 2009; Roland & Roland, 1980).

2.2. Morphological adaptations: shape and size of organs

The external appearance of hydrophytes reflects “economy” on mechanical tissues and the desire to increase the surface area of contact with water:

  • Leaves of submerged plants are usually thin, delicate, often repeatedly dissected into thread-like lobes. This shape dramatically increases the area of contact with water, facilitating the absorption of carbon dioxide and minerals, and reduces water resistance during movement (Strasburger et al., 1971). For example, the leaves of Myriophyllum resemble feathery “fir trees”, and in Ranunculus aquatilis the submerged leaves are strongly dissected, while the emergent ones are entire or lobed (heterophylly).

  • Stems of hydrophytes are long, thin, and easily bend. In free-floating forms, they are often shortened or reduced (e.g., in duckweed Lemna the stem is transformed into a small green leaf-like cladode).

  • Root system in many hydrophytes is poorly developed, root hairs are absent or few, as the absorption of water and salts occurs through the surface of stems and leaves. In some submerged species (e.g., Ceratophyllum) roots are completely absent — the plant absorbs nutrients directly through epidermal cells (Roland & Roland, 1980).

2.3. Anatomical adaptations: tissues, conducting elements and intercellular spaces

The internal structure of hydrophytes is a classic example of hydromorphosis, i.e., a complex of traits opposite to xeromorphic ones:

  1. Epidermis and cuticle: in submerged hydrophytes, epidermal cells are thin-walled, the cuticle is either absent or very thin and easily permeable to water and gases. In floating leaves, the cuticle is developed on the upper side (facing the air), and stomata are located only on the upper side (epistomatic type) (Evert, 2006). In submerged leaves, stomata are usually absent.

  2. Mesophyll: in submerged leaves, differentiation into palisade and spongy parenchyma is often absent. Mesophyll cells are large, with large chloroplasts, loosely arranged. In floating leaves, palisade parenchyma may be well developed, but beneath it lies a powerful aerenchyma (air-bearing tissue).

  3. Aerenchyma — the key tissue of hydrophytes. It is a parenchyma in which intercellular spaces merge into large air cavities separated by thin partitions (Roland & Roland, 1980; Beck, 2010). Aerenchyma performs three functions:

    • provides buoyancy and keeps leaves at the water surface;

    • serves as a reservoir of oxygen coming from the aerial parts or released during photosynthesis;

    • through the air cavities, oxygen diffuses to the roots, preventing their acidification in anaerobic bottom sediments.

The volume of aerenchyma can reach 70% or more of the total organ volume. For example, in the stems of Elodea, intercellular spaces are already visible under low magnification (Yakovlev et al., 2008).

  1. Conducting and mechanical tissues: xylem (water-conducting vessels) is reduced in hydrophytes, since water and salts are absorbed over the entire surface and move mainly through the apoplast of parenchyma cells. Vessels often have thin walls, and lignin is deposited in small amounts. Mechanical tissues (collenchyma, sclerenchyma) are poorly developed or absent, as water itself supports the plant. Consequently, hydrophytes tear easily when taken out of water and quickly wither when removed (Strasburger et al., 1971).

2.4. Features of reproduction and dispersal

The aquatic environment leaves its mark on the reproductive biology of hydrophytes:

  • Vegetative reproduction often predominates (by shoot fragmentation, brood buds, tubers). For example, Canadian waterweed (Elodea canadensis) reproduces exclusively vegetatively in Europe, as it was introduced only in the female form (Scremin-Dias, 2009).

  • Flowers of many hydrophytes rise above the water (or bloom on the surface) and are pollinated by insects or wind. In some species (e.g., Vallisneria and Elodea), hydrophily — underwater pollination where pollen is carried by water currents — has developed.

  • Fruits and seeds have adaptations for dispersal by water (hydrochory): spongy tissue, air sacs, non-wettable coats.

2.5. Ecological and applied significance of hydrophytes

Hydrophytes play a key role in aquatic ecosystems: they serve as shelter for fish and invertebrates, participate in water self-purification (accumulate heavy metals and radionuclides), and are a food base for waterfowl and mammals. However, when they grow massively (e.g., Eichhornia crassipes in the tropics, Elodea canadensis in temperate latitudes), they can become weeds, hindering navigation and fishing (Scremin-Dias, 2009).

In applied terms, the study of hydrophytes is important for:

  • biological wastewater treatment (design of “planted treatment facilities” using aquatic plants);

  • landscape design (ornamental ponds);

  • understanding the early stages of land plant evolution, as hydrophytes retain many primitive structural features (absence of cuticle, weak xylem development, simple life cycles) (Beck, 2010).

In the next section, we will consider the intermediate group — mesophytes, which are the most typical for most cultivated plants and serve as a kind of “reference” for comparison.

3. Group 2: Hygrophytes (Hygrophytes)

Hygrophytes (from Greek hygros — wet and phyton — plant) are an ecological group of plants living in conditions of high air and soil moisture, but not submerged in water. Unlike hydrophytes, hygrophytes are terrestrial plants, but their habitats are characterized by constantly high substrate moisture (banks of water bodies, wet meadows, swamps, tropical rainforests) or air saturated with water vapour (fog forests, gorges, caves) (Yakovlev et al., 2008; Serebryakova et al., 2006).

Unlike xerophytes, which conserve water, and hydrophytes, which are submerged in it, hygrophytes face a paradoxical situation: with an abundance of water in the environment, they experience difficulties with transport and gas exchange, because high air humidity greatly reduces the water vapour gradient between the leaf and the atmosphere. This leads to a slowdown in transpiration as the driving force of root pressure, as well as a risk of liquid wetting of the leaf surface, which hinders normal aeration (Evert, 2006).

3.1. Ecological conditions of hygrophyte habitats

Hygrophytes include plants growing in the following biotopes:

  • River floodplains and riparian zones (e.g., marsh marigold Caltha palustris, purple loosestrife Lythrum salicaria, water forget-me-not Myosotis scorpioides).

  • Wet and swampy forests (e.g., touch-me-not balsam Impatiens noli-tangere, stinging nettle Urtica dioica, wood sorrel Oxalis acetosella, many ferns).

  • Tropical rainforests (hylea), where relative air humidity approaches 100% and epiphytes and terrestrial herbs with thin, delicate leaves abound (Beck, 2010).

A characteristic feature of hygrophytes is that they cannot tolerate even short-term drying of the substrate or a decrease in air humidity. When water is lacking, their leaves quickly lose turgor, wilt and die, as their anatomy is not adapted to reduce transpiration.

3.2. Morphological features of hygrophytes

The external appearance of hygrophytes reflects “adaptation to excess moisture” and often to shading (since wet forests are usually densely shaded):

  1. Leaf blades are large, thin, delicate, often with a smooth, shiny surface. In many forest hygrophytes (e.g., false lily of the valley Maianthemum bifolium), the leaves have a characteristic “watery” consistency and are easily damaged by mechanical impact.

  2. Colour is bright green due to high chlorophyll content and the absence of protective waxy coatings and pubescence (Sorokopudov et al., 2018 — in context of R. rugosa, comparing pubescence with hygrophytes).

  3. Root system is superficial, often with adventitious roots, may be poorly developed as water is easily accessible. Some hygrophytes form additional adventitious roots on stems (e.g., hedge bindweed Calystegia sepium).

  4. Stems are succulent, soft, often hollow (with aerenchyma) or with a loose pith, which promotes better gas exchange and facilitates water transport. In many forest herbs, stems are erect but easily lodge.

  5. Pubescence in hygrophytes is absent or represented by sparse, delicate, unicellular hairs that do not hinder evaporation but may instead help retain water droplets on the surface (which is sometimes undesirable as it promotes fungal diseases) (Evert, 2006).

3.3. Anatomical adaptations of the hygrophyte leaf

The internal structure of the hygrophyte leaf most clearly demonstrates signs of hygromorphosis:

  1. Epidermis:

    • Epidermal cells are large, thin-walled, with a very thin cuticle or none. This facilitates the absorption of water and gases directly through the leaf surface.

    • Outer cell walls are often convex (papillose), which increases the surface for condensation of water vapour from the air (Roland & Roland, 1980).

  2. Stomata:

    • Located on both sides of the leaf (amphistomatic type) or only on the lower side (hypostomatic), but their density (number per 1 mm2) is usually low compared to xerophytes (Sorokopudov et al., 2018 — in R. rugosa under high humidity, stomatal number is lower).

    • Stomata are not sunken, often protrude above the epidermal surface. Guard cells have thin walls, chloroplasts are well developed, and the opening mechanism triggers at the slightest increase in turgor.

    • Under saturated humidity, stomata may be constantly open, even at night.

  3. Mesophyll:

    • Clearly differentiated into palisade (usually single-layered, cells not very densely packed) and spongy layers. Spongy parenchyma occupies most of the leaf volume, contains very large intercellular spaces (aerenchyma) that connect with stomata through substomatal cavities (Evert, 2006; Beck, 2010).

    • Thanks to the well-developed system of intercellular spaces, oxygen and carbon dioxide diffuse rapidly to the photosynthesising cells even when gas exchange is hindered (e.g., in water droplets on leaves).

  4. Hydathodes:

    • At the ends of veins or along leaf margins in many hygrophytes (e.g., marsh marigold, loosestrife, some grasses), hydathodes are located — special water stomata or groups of cells (epithem) through which water droplets are actively secreted (guttation) (Evert, 2006).

    • Guttation occurs during night and morning hours, when root pressure is high and transpiration is low due to air humidity. Thus, the plant gets rid of excess water taken up by the roots.

  5. Conducting and mechanical tissues:

    • Xylem is moderately developed, vessels are wide but do not always have thick lignified walls. Water flow is slow because the water potential gradient is small.

    • Mechanical tissues (sclerenchyma, collenchyma) are weakly expressed, which accounts for the softness and delicacy of the shoots.

3.4. Water regime and physiological features

The water exchange of hygrophytes is characterized by high transpiration intensity with low water-holding capacity of tissues (Yakovlev et al., 2008). Main indicators:

  • Water deficit in tissues is usually small (about 5–10% of saturation) due to constant water supply.

  • Tissue hydration is high (total water content can reach 85–90% of fresh weight).

  • When water supply is disrupted (e.g., during transplantation or short-term drought), hygrophytes quickly lose turgor and are irreversibly damaged, because their cells are not able to actively synthesise osmoprotectants (proline, sugars) and retain water (Sorokopudov et al., 2018).

3.5. Applied significance and indicator role

Hygrophytes have important economic significance and are used for scientific purposes:

  1. Indicators of soil water regime:

    • The presence and abundance of hygrophytes (marsh marigold, marsh horsetail Equisetum palustre, marsh cinquefoil Comarum palustre) indicates a shallow water table or excessive moisture.

    • In agriculture, such areas are considered “problematic”: they require drainage for growing mesophytic crops.

  2. Land reclamation:

    • Hygrophytes (e.g., willows Salix spp., alder Alnus spp., some sedges) are often planted along the banks of water bodies to stabilise them and regulate water regime.

  3. Ornamental horticulture:

    • Many hygrophytes (daylilies, swamp irises, marsh marigolds, hostas) are used to design the shoreline of artificial ponds, wet flower beds (“bogs”) and container water gardens.

  4. Biological purification:

    • Hygrophytes, together with hydrophytes, are used in phytoremediation systems to absorb excess nutrients (nitrogen, phosphorus) from wastewater, as they actively accumulate mineral substances in their aboveground organs (Scremin-Dias, 2009).

In the next section, we will consider mesophytes — plants with a balanced water regime, to which most agricultural crops and many temperate zone plants belong.

4. Group 3: Mesophytes (Mesophytes)

Mesophytes (from Greek mesos — middle, intermediate and phyton — plant) are an ecological group of plants adapted to living in conditions of moderate, balanced moisture. This is the most extensive and diverse group, to which the vast majority of agricultural crops belong (wheat, maize, soybean, potato, tomatoes, rice — although rice is often classified as a hydrophyte or hygrophyte by growing conditions, botanically it is a mesophyte), as well as many meadow and forest herbs and deciduous trees of the temperate zone (Yakovlev et al., 2008; Serebryakova et al., 2006).

Mesophytes occupy an intermediate position between hygrophytes and xerophytes, so their anatomical and morphological features are considered “baseline” or “typical” for terrestrial flowering plants. They do not have pronounced adaptations either to drought or to waterlogging, but at the same time possess sufficient plasticity to withstand short-term deviations of water regime from the optimum (Beck, 2010).

4.1. Ecological conditions and distribution

Mesophytes dominate in climatic zones with uniform rainfall distribution and moderate temperatures — in broad-leaved and mixed forests, floodplain and upland meadows, and agrocenosis fields. The optimal soil moisture for them is 60–80% of full water capacity, and relative air humidity is 60–80% (Evert, 2006).

Compared to hygrophytes, mesophytes are more tolerant of short-term moisture reduction (drought), and compared to xerophytes, more tolerant of temporary increases (flooding). However, prolonged exposure to extreme conditions (several weeks of drought or waterlogging) leads to growth inhibition, reduced yield, and often death.

4.2. Morphological features of mesophytes

The external appearance of mesophytes can be considered “typical” for the idea of a flowering plant:

  • Leaf blade of moderate size, medium thickness, usually green, without waxy coating or with a thin coating, usually smooth, rarely slightly pubescent. Leaf shape varies from broadly ovate to lanceolate, margin may be entire, toothed or serrated (Beck, 2010).

  • Stem erect, elastic, with well-expressed mechanical tissues, but not woody (in herbs) or moderately woody (in trees and shrubs). Pith is loose, but without large air cavities (as in hydrophytes).

  • Root system is well developed, usually taproot (in dicots) or fibrous (in monocots), with many root hairs providing efficient water uptake from the soil.

4.3. Anatomical features (using the leaf as an example)

The mesophyte leaf is a convenient object for studying typical anatomical structure (Evert, 2006; Roland & Roland, 1980):

  1. Epidermis: single-layered, cells tightly appressed, outer walls moderately thickened, covered with a thin but continuous cuticle (without heavy wax deposits). The cuticle protects against excessive evaporation but does not hinder normal gas exchange.

  2. Stomata:

    • Located mainly on the lower side of the leaf (hypostomatic type), less often on both sides (amphistomatic).

    • Stomatal density ranges from 100 to 300 per 1 mm2 (average values).

    • Stomata are at the level of epidermal cells or slightly raised, have well-developed bean-shaped guard cells capable of actively regulating opening depending on turgor and signals (abscisic acid, light) (Chen et al., 2024 — in the context of ABA signalling).

    • The dynamics of opening/closing are fast (from a few minutes to an hour), allowing the plant to finely regulate transpiration during the day.

  3. Mesophyll:

    • Clearly differentiated into palisade (columnar) and spongy parenchyma.

    • The palisade layer (1–2 layers of cylindrical cells) is located under the upper epidermis; the cells contain many chloroplasts and are oriented perpendicular to the surface (maximising light capture).

    • Spongy parenchyma occupies the lower part of the leaf, cells are rounded, with large intercellular spaces ensuring gas circulation (CO2 and O2).

    • The ratio of palisade to spongy parenchyma is approximately 1:1 by volume.

  4. Vascular bundles (veins):

    • Of the collateral type, with xylem (water-conducting tissue) facing the upper side of the leaf and phloem (organic substances) facing the lower side.

    • Xylem contains both tracheids and vessels (in dicots), walls are thickened and lignified, but not excessively.

    • Bundles are surrounded by a parenchymatous sheath, sometimes with starch-storing cells (Beck, 2010).

  5. Mechanical tissues: collenchyma and sclerenchyma are moderately expressed, located along the major veins and at the leaf margin, providing elasticity but not rigidity.

4.4. Water regime and physiological parameters

Mesophytes are characterized by average values of water exchange, which can shift temporarily with changing conditions:

  • Total tissue hydration is 70–80% (Yakovlev et al., 2008).

  • Water deficit normally does not exceed 10–15%; during drought it can increase to 25–30% without irreversible damage (thanks to the ability for osmotic regulation — accumulation of proline and sugars).

  • Transpiration is intensive (a plant can evaporate an amount of water equal to its own mass per day), creating sufficient force for water ascent from roots to leaves.

  • Stomatal regulation is effective: under mild water deficit, stomata partially close, reducing transpiration; when water supply is restored, they open again.

4.5. Mesophytes in agronomy: importance and vulnerability

Most crop plants that form the basis of global food production are mesophytes. Their high productivity (yield) is achieved only by maintaining an optimal water regime through irrigation and agricultural practices. In dry years or when water supply is disrupted, mesophytes sharply reduce yields due to:

  • stomatal closure and reduced photosynthesis;

  • disruption of assimilate transport;

  • oxidative stress in chloroplasts (accumulation of reactive oxygen species due to excess light when stomata are closed) (Chen et al., 2024).

For this reason, breeders and physiologists are seeking ways to increase drought tolerance of mesophytes using genetic material from xerophytes (wild relatives) (Chen et al., 2024). For example, introducing the SUB1A gene from xerophytic rice into cultivated varieties has increased their tolerance to submergence (Iftikhar et al., 2025). Similarly, genes controlling stomatal development (EPF, TMM) can be used to optimise water use in mesophytes.

Thus, mesophytes are the most important group for the agronomist and ecologist, as they form the basis of anthropogenic ecosystems (agrocenoses) and are most sensitive to climate change. Understanding their “baseline” structure is necessary for comparison with xerophytes (drought-tolerant) and hygrophytes (moisture-loving), to which we turn in the next section.

5. Group 4: Xerophytes (Xerophytes)

Cross sections of a mesophyte and xerophyte leaf
©Bgqhrsnogcommons.wikimedia.orgCC BY-SA 3.0

Cross section of a xerophyte leaf

The diagram demonstrates key xeromorphic adaptations: multilayered palisade parenchyma, thick cuticle and sclerenchyma strands. C – thick cuticle, E – multilayered epidermis, H – dead epidermal hairs, P – multilayered palisade and spongy parenchyma, S – internal stomata.

Xerophytes (from Greek xeros — dry and phyton — plant) are an ecological group of plants living under conditions of chronic or seasonal deficit of available water. These can be arid steppes, semi-deserts and deserts, rocky slopes and cliffs, as well as drying sandbanks. A special form of “physiological drought” (when water is present but unavailable due to high osmotic pressure of the soil solution) is experienced by halophytes (salt marsh plants), which often exhibit xeromorphic features (Yakovlev et al., 2008; Serebryakova et al., 2006).

Unlike mesophytes, which only temporarily tolerate water shortage, xerophytes possess a complex of inherited adaptations that allow them to successfully vegetate and reproduce under negative water balance (transpiration exceeds water uptake over a long period). Two main strategies for survival in arid conditions are distinguished (Chen et al., 2024; Beck, 2010):

  1. Drought avoidance — plants complete their life cycle (flower and fruit) in a short wet period (ephemerals and ephemeroids) or have deep root systems that allow them to extract water from lower soil horizons.

  2. Drought tolerance — plants actively regulate water balance by reducing transpiration and/or storing water in tissues. Among tolerant xerophytes, two subgroups are distinguished:

    • Succulents (from Latin succulentus — juicy) — store water in leaves, stems or roots.

    • Sclerophytes (from Greek skleros — hard, rigid) — limit transpiration and withstand tissue dehydration thanks to powerful mechanical elements and dense structure.

5.1. Succulents — living water “reservoirs”

Succulents accumulate moisture in specialised water-storage parenchyma, which can occupy up to 90% of the organ’s volume. Based on the location of the water-storage tissue, the following are distinguished:

  • Leaf succulents — water is stored in thickened, often fleshy leaves (aloe Aloe spp., agave Agave spp., stonecrops Sedum spp., houseleeks Sempervivum). In some leaf succulents (e.g., Lithops — “living stones”), the leaves are transformed into two almost fused fleshy lobes, and photosynthesis occurs through special “windows” on their surface (Evert, 2006).

  • Stem succulents — water is stored in a thickened stem, and leaves are reduced to spines or small scales (cacti Cactaceae, many euphorbias Euphorbia from African deserts, stapeliads Stapelia). The ribbed shape of the stem (in cacti) allows the plant to shrink when water is lost and expand when it is stored, without damaging tissues (Serebryakova et al., 2006).

  • Root succulents — storage roots (e.g., in some Ceropegia, Pelargonium).

The anatomy of succulents is characterised by the following features:

  • Powerful water-storage parenchyma with large vacuolated cells containing mucous substances that retain water.

  • Epidermal cells are often covered with a thick cuticle and waxy layer, stomata are sunken and open mainly at night (CAM photosynthesis), which sharply reduces water loss during the day (Evert, 2006; Chen et al., 2024).

  • The chlorophyll-bearing layer (chlorenchyma) is located directly under the epidermis, even with significant thickening of the stem or leaf, ensuring photosynthesis with minimal surface area.

  • Mechanical tissues are poorly developed, as the supporting function is partly performed by the turgor of water-filled cells.

5.2. Sclerophytes — water conservation through “rigidity”

Sclerophytes (from Greek skleros — hard) do not store water, but actively prevent its evaporation and withstand strong tissue dehydration (up to 30–40% of the original water content) without loss of viability. Typical representatives: feather grasses Stipa spp., sheep fescue Festuca valesiaca, saxaul Haloxylon spp., oleander Nerium oleander, olive Olea europaea, many eucalypts Eucalyptus and acacias Acacia (especially those with phyllodes) (Yakovlev et al., 2008; Sorokopudov et al., 2018 — in R. rugosa under arid conditions, scleromorphic features appeared).

Sclerophytes exhibit a complex of xeromorphic traits, many of which are directly opposite to hydromorphic ones:

  1. Leaves: small, narrow, hard (leathery), often with downward-rolled margins (in grasses) or rolled into a tube. In some sclerophytes, leaves are completely reduced, and green stems (phylloclades, cladodes) carry out photosynthesis (Beck, 2010). In eucalypts, leaf blades are turned edge-on to the sun, reducing heating and evaporation.

  2. Epidermis:

    • Multilayered (2–4 layers), cells with strongly thickened outer and lateral walls, covered with a thick cuticle and often a waxy layer (in some species, so much wax is produced that it is harvested — carnauba palm).

    • Dense pubescence (trichomes) — reflection of sunlight and creation of a “cushion” of humid air near the leaf surface. In oleander, hairs line the stomatal crypts (depressions on the lower leaf surface) (Evert, 2006).

  3. Stomata:

    • Number reduced (sometimes to 50–100 per 1 mm2 compared to 200–300 in mesophytes), sunken in crypts, often covered with hairs, which significantly reduces transpiration even when stomata are open (Sorokopudov et al., 2018 — in R. rugosa under arid conditions, stomatal number decreased).

    • Guard cells have thickened walls; some sclerophytes can completely plug the stomatal slits with wax “plugs”.

  4. Mesophyll:

    • Palisade parenchyma is multilayered (3–5 layers), cells compactly packed, often folded (in pines — “folded chlorenchyma”), which increases the surface area of chloroplasts within a limited volume (Evert, 2006).

    • Spongy parenchyma is almost absent, intercellular spaces are minimised — this reduces the risk of excessive evaporation but requires more active gas exchange through stomata.

  5. Conducting and mechanical tissues:

    • Xylem is powerfully developed, vessels with thick lignified walls, often with libriform fibres.

    • Sclerenchyma (often as a continuous ring under the epidermis or around vascular bundles) provides mechanical strength even under severe dehydration, when parenchyma loses turgor.

5.3. Root system adaptations of xerophytes

Water supply to xerophytes is as important as reducing losses. Root systems can be of two types:

  • Deep taproots (in camel thorn Alhagi maurorum, saxaul) reach groundwater at depths of 10–20 m or more, providing the plant with moisture throughout the dry season (Yakovlev et al., 2008).

  • Extensive superficial root systems (in cacti, some ephemerals) are branched in the upper (0–50 cm) soil layer and can quickly absorb water after rare rains, even with small amounts of precipitation.

Anatomically, roots of xerophytes often develop a multilayered exodermis with suberised cells, reducing water loss from the cortex, as well as a powerful central cylinder with a large number of xylem elements (Evert, 2006).

5.4. Physiological features and significance for breeding

Xerophytes possess unique physiological mechanisms that are increasingly becoming targets of biotechnological research to improve drought tolerance of crop plants:

  • Low basal level of abscisic acid (ABA) and altered structure of ABA receptors (PYR/PYL), providing a fast and sensitive stomatal response to water deficit (Chen et al., 2024).

  • Efficient mechanism for scavenging and neutralising reactive oxygen species (high activity of superoxide dismutase, catalase, peroxidases) (Sorokopudov et al., 2018).

  • CAM photosynthesis in leaf and stem succulents: stomata open at night, CO2 is fixed into organic acids (malic acid), which are decarboxylated during the day, and CO2 is used in the Calvin cycle with stomata closed. This reduces water loss tens of times compared to C3 plants (Evert, 2006).

In recent years, the genomes of xerophytes (e.g., Populus euphratica, Zygophyllum xanthoxylum) have been actively studied to identify genes responsible for salt and drought tolerance. Genes NHX (vacuolar Na+/H\+ antiporters), AVP1 (H\+-pyrophosphatase), ERF-VII and SnRK2 are already being used to create transgenic mesophytes (soybean, rice, wheat) with enhanced drought tolerance (Chen et al., 2024; Iftikhar et al., 2025).

5.5. Ecological and economic role of xerophytes

Xerophytes form the plant cover of arid and semi-arid zones of the Earth, preventing soil erosion and desertification. Many of them are valuable forage (camel thorn, feather grass, wormwood), medicinal (aloe, liquorice), food (date palm, prickly pear) or ornamental (cacti, stonecrops, houseleeks) plants. Studying their adaptations is necessary not only for fundamental biology but also for solving practical problems: breeding drought-tolerant varieties of agricultural crops and restoring degraded lands (Chen et al., 2024).

Next, in the “Applied significance” section, we will consider how knowledge of adaptations to water regime is used in agronomy, breeding and environmental conservation.

6. Comparative table: morphological and anatomical adaptations to water regime

For a visual summary of the material presented in the previous sections, a summary table is given below. It compares the key features of vegetative organs (leaf, stem, root) for four main ecological groups: hydrophytes, hygrophytes, mesophytes and two subgroups of xerophytes — succulents and sclerophytes. Such a table is an essential element of educational courses on ecological plant morphology at universities in the USA, Germany and the Netherlands (Beck, 2010; Evert, 2006; Raunkiaer, 1934).

Trait Hydrophytes (aquatic) Hygrophytes (moisture-loving) Mesophytes (moderate) Xerophytes (drought-tolerant)
Succulents Sclerophytes
Cuticle and waxy layer Absent or very thin, easily permeable to water and gases Thin, does not hinder evaporation Moderate thickness, continuous, with a slight waxy layer Thick, shiny, often with a heavy wax layer (sometimes up to 5 mm) Very thick, often multilayered, with wax deposits
Stomata (location, density, depth) Absent in submerged forms; in floating leaves only on upper side (epistomatic). Density low or zero On both sides (amphistomatic), often raised above the surface, density low (50–150/mm2) Mainly on lower side (hypostomatic), density medium (100–300/mm^2^), at epidermis level Few (rare), open at night (CAM photosynthesis), often sunken in crypts Few (50–100/mm2), deeply sunken in crypts, often covered by hairs or wax plugs
Epidermal cells Large, thin-walled, often contain chloroplasts Large, thin-walled, with convex outer walls (papillae) Medium-sized, tightly appressed, outer walls moderately thickened Large, thin-walled (water-storing) or multilayered hypodermis Small, with strongly thickened outer and lateral walls, often multilayered hypodermis
Mesophyll Poorly differentiated (no palisade), cells large, loose; intercellular volume >50% Differentiated, palisade layer single-layered, spongy parenchyma with large intercellular spaces Clearly differentiated (1–2 layers of palisade, spongy parenchyma with intercellular spaces 20–30% of volume) Chlorenchyma under epidermis (sometimes folded), central part occupied by large water-storage cells Multilayered palisade parenchyma (3–5 layers), compact, spongy parenchyma almost absent
Aerenchyma (air-bearing tissue) Well developed in stems and leaves (up to 70% of volume), large cavities Developed in stems and petioles, especially in marsh forms Poorly developed, only in loose pith of stem Absent (replaced by water-storage parenchyma) Absent or reduced (very small intercellular spaces)
Mechanical tissues (collenchyma, sclerenchyma) Almost absent (water supports the body) Weakly developed, stems and leaves soft Moderately developed (provide elasticity) Weakly developed (turgor serves as support) Strongly developed (sclerenchyma fibres, rings, strands), tissues dry and rigid
Xylem (water-conducting tissue) Reduced, vessels with thin walls, often absent Poorly developed, vessel walls not always lignified Well developed, vessels with thickened lignified walls Moderately developed, but water-storage parenchyma predominates Powerfully developed, large vessels, many libriform fibres
Root system Weak, root hairs absent or few, often no roots Superficial, often with adventitious roots on stems Taproot or fibrous, with dense root hairs Superficial, widely branched (catching dew) or taproot, deeply penetrating (up to 20 m) Deep taproot or highly branched superficial with powerful root hairs
Special structures Hydathodes (in floating forms), leuco- and chloroplasts in epidermis, heterophylly Hydathodes at leaf margins (guttation), delicate trichomes, often absence of pubescence No pronounced special structures; pubescence weak Water-storage parenchyma in leaves, stems or roots; CAM photosynthesis Stomatal crypts, dense pubescence (sometimes felt-like), folded chlorenchyma, wax “plugs”
Example genera Elodea, Ceratophyllum, Nymphaea, Lemna, Vallisneria Caltha, Impatiens, Urtica, Oxalis, Myosotis, Pulmonaria Triticum, Zea, Solanum, Brassica, Quercus, Betula, Rosa Aloe, Sedum, Sempervivum, Lithops, Opuntia, Euphorbia (succulent) Stipa, Festuca, Haloxylon, Nerium, Olea, Pinus, Calligonum

Explanatory notes for the table:

  • Hydrophytes and hygrophytes are similar in the abundance of aerenchyma and thin integuments, but hydrophytes are more reduced due to submergence in water.

  • Mesophytes — the “golden mean”: they have no extreme adaptations but possess all the necessary tissues for active water exchange.

  • Xerophytes are divided into two strategies: succulents (water storage, CAM photosynthesis) and sclerophytes (rigidity, strong reduction of transpiration). Many xerophytes combine elements of both subgroups (e.g., leaf succulents often have a scleroified hypodermis).

This table can serve as a reference outline for students and as a basis for laboratory work on microscopic analysis of leaf anatomy in plants from different ecological groups.

In the next section (7. Applied significance), we will discuss how knowledge of these adaptations is used in agronomy, breeding, land reclamation and nature conservation.

7. Applied significance for the agricultural sector

The study of morphological and anatomical adaptations of plants to water regime has not only theoretical but also great practical importance for agriculture and forestry, breeding, land reclamation, landscape design and nature conservation. Knowledge of ecological groups allows the agronomist and plant grower to predict crop behaviour on different soils, select optimal irrigation regimes, conduct targeted breeding for drought or salt tolerance, and use indicator plants to assess land condition (Yakovlev et al., 2008; Serebryakova et al., 2006). The main directions of applied use of this knowledge are listed below.

7.1. Indication of soil and hydrological conditions (phytoremediation)

Plants respond sensitively to the water regime of the substrate, so their presence, abundance and condition can serve as a reliable indicator of soil moisture, depth of groundwater and degree of salinity. This is especially important for agricultural land use and land reclamation.

  • Hygrophytes as indicators of waterlogging: mass occurrence of marsh marigold (Caltha palustris), purple loosestrife (Lythrum salicaria), meadowsweet (Filipendula ulmaria), marsh horsetail (Equisetum palustre) and some sedges (Carex spp.) indicates shallow groundwater (less than 0.5–1 m), high soil moisture and the need for drainage to grow most crops (Yakovlev et al., 2008).

  • Mesophytes as indicators of normal moisture: the presence of typical meadow and forest species (red clover Trifolium pratense, timothy Phleum pratense, meadow foxtail Alopecurus pratensis) indicates a water regime optimal for field crops (60–80% of full water capacity).

  • Xerophytes and halophytes as indicators of water deficit and salinity: dominance of feather grasses (Stipa spp.), wormwoods (Artemisia spp.), prostrate kochia (Kochia prostrata) and saltworts (Salsola spp.) indicates low water availability and often soil salinity. Some species (glasswort Salicornia europaea, seepweed Suaeda) are indicators of severe salinity (Yakovlev et al., 2008; Chen et al., 2024).

In practice, when assessing soil quality and planning reclamation measures, vegetation maps are drawn up with contours occupied by different ecological groups. This allows assessment of the hydrological regime of an area without expensive drilling.

7.2. Breeding for drought and salt tolerance

Xerophytes (including wild relatives of crop plants and halophytes) are a valuable genetic reservoir of traits that can be introgressed into mesophytic varieties to increase their resistance to abiotic stresses (Chen et al., 2024; Iftikhar et al., 2025).

The main targets for breeding, identified from the study of xerophytes, are:

  • Stomatal traits: reduced stomatal density (genes EPF1/2, TMM), sunken stomata in crypts, presence of pubescence, rapid stomatal closure under water deficit (controlled by ABA signalling — receptors PYR/PYL, protein phosphatases PP2C, kinases SnRK2/OST1). Transgenic lines of rice, wheat and barley with reduced stomatal density have already been created, which save water without significant yield loss (Chen et al., 2024).

  • Water storage traits: genes of vacuolar Na+/H\+ antiporters (NHX1) and H\+-pyrophosphatase (AVP1) that ensure ion accumulation in vacuoles and increased turgor at low water potential. Transgenic plants with increased expression of these genes (tomatoes, rapeseed, rice, cotton) have shown increased drought and salt tolerance (Chen et al., 2024; Iftikhar et al., 2025).

  • Membrane stabilisation and antioxidant defence traits: genes encoding superoxide dismutase (SOD), catalase (CAT), peroxidases, as well as enzymes of proline and trehalose synthesis. These genes are often overexpressed in xerophytes and halophytes (Chen et al., 2024). For example, the TPP7 gene (trehalose-6-phosphate phosphatase) in rice, borrowed from xerophytic forms, provides tolerance to submergence (anaerobic stress) (Iftikhar et al., 2025).

  • CAM photosynthesis: although transferring the CAM pathway into C3 crops is technically challenging, work is underway to use genes of phosphoenolpyruvate carboxylase (PEPC) and malate dehydrogenase from succulents (cacti, agaves) to increase water use efficiency of crops (Evert, 2006).

A classic example of successful breeding is the introduction of the SUB1A gene (from xerophytic rice tolerant to submergence) into high-yielding rice varieties, allowing them to withstand complete flooding for up to two weeks (Iftikhar et al., 2025).

7.3. Land reclamation and water management

Understanding the water regime of plants underlies the design of drainage and irrigation systems.

  • Drainage reclamation (drainage): on waterlogged lands occupied by hygrophytes, installation of drainage lowers the groundwater level, which gradually leads to a replacement of hygrophytes by mesophytes and makes the soil suitable for agricultural use. At the same time, some hygrophytes (willows, alder, poplar) are themselves used for biodrainage — they are planted along field edges to actively pump water out of the soil (transpiration “pump”).

  • Irrigation reclamation: during irrigation, it is necessary to take into account that xerophytes, even with excess water, retain xeromorphic features (thick cuticle, sunken stomata), which can lead to leaf overheating on hot days because evaporation is hindered. Therefore, the irrigation regime (frequency and amount of watering) for drought-tolerant varieties should be special (less often but more abundantly), unlike for mesophytes.

  • Phytoremediation of saline soils: halophytes (salt-tolerant xerophytes) are used for salt extraction — they absorb salts from the soil and accumulate them in aboveground organs, which are then removed. This method is used in Central Asian and Middle Eastern countries (Chen et al., 2024).

7.4. Landscape design and ornamental horticulture

When creating gardens, parks and green areas, it is important to select plants according to the natural moisture of the site in order to avoid their death and excessive watering.

  • For wet, swampy areas (pond banks, lowlands), hygrophytes and hydrophytes are used: hostas, swamp iris (Iris pseudacorus), marsh marigold, water arum (Calla palustris), water lilies, yellow water lilies, water hyacinth (in containers) (Scremin-Dias, 2009).

  • For dry, rocky areas (rock gardens, rockeries, xeric gardens), succulents (stonecrops, houseleeks, jade plants) and sclerophytes (feather grasses, wormwoods, lavender, rosemary, yucca, cacti) are used. In arid regions, xeriscaping (from English xeriscaping) — landscape design with minimal water use based on xerophytes — is becoming increasingly widespread (Raunkiaer, 1934; Niklas, 2008).

7.5. Nature conservation and restoration of disturbed lands

  • Biodiversity conservation: knowledge of ecological groups is necessary for the creation of protected areas (nature reserves, national parks), since hydrophytes and hygrophytes are often rare and vulnerable species that require maintenance of the natural hydrological regime.

  • Reclamation of technogenically disturbed lands (quarries, dumps, deserts) is carried out using pioneer xerophytes (psammophytes — plants of sands) that stabilise loose substrates and create conditions for subsequent vegetation restoration. For example, sand acacia (Ammodendron) and calligonum (Calligonum) are used to fix dune sands (Yakovlev et al., 2008; Beck, 2010).

Thus, knowledge of morphological and anatomical adaptations of plants to water regime is an integral part of the training of agronomists, ecologists and landscape designers. It allows rational use of water resources, increasing the resilience of agroecosystems to droughts and floods, and preserving natural diversity in a changing climate.

References

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