Plant cell
Plant cell — is a structural, functional, and genetic unit of a plant organism, representing a complex eukaryotic system characterized by the presence of a cell wall, plastids, and a large central vacuole (Yakovlev et al., 2001; Evert, 2006). Unlike animal and fungal cells, the plant cell is capable of autotrophic nutrition due to its photosynthetic organelles — chloroplasts, and its shape and mechanical strength are maintained by the cellulose wall. The vacuole, occupying up to 90% of the volume of a mature cell, performs the functions of osmotic regulation, storage of metabolites, and deposition of end products of metabolism (Andreev, 2001). The totality of all cells connected by plasmodesmata forms a single symplast, emphasizing the integrative nature of the plant organism (Evert, 2006).
Understanding the structure and vital activity of the plant cell is of fundamental importance for agricultural sciences. It is at the cellular level that key processes determining the productivity of agricultural crops are realized: photosynthesis, mineral nutrition, assimilate transport, growth, and resistance to biotic and abiotic stresses (Mauseth, 2017). Knowledge of the mechanisms of the cell cycle and differentiation allows breeders and biotechnologists to control organogenesis, achieving increased biomass, altered plant architecture, and improved fruit and seed quality (Francis, 2007; Lee and Torii, 2026). Methods of in vitro cell and tissue culture, based on the phenomenon of totipotency, are widely used for vegetative propagation of valuable genotypes, obtaining virus-free planting material, and accelerating the breeding process (Fehér, 2019).
Furthermore, research on the plant cell underpins modern biotechnologies aimed at increasing plant stress tolerance. The study of vacuolar transporters and xenobiotic detoxification systems opens pathways for creating varieties tolerant to soil salinity and heavy metal pollution (Andreev, 2001). Understanding the molecular basis of cellular responses to pathogens and herbicides contributes to the development of environmentally friendly methods of plant protection (Mauseth, 2017). Thus, plant cytology serves as a theoretical foundation for solving applied problems in agronomy, from increasing yields to sustainable agriculture.
1. Key differences between plant and animal cells
Despite the general eukaryotic plan of organization, cells of higher plants exhibit a number of fundamental differences from animal cells, reflecting the features of their autotrophic lifestyle and fixed position in space (Serebryakova et al., 2006; Graham et al., 2014).
The first and most obvious difference is the presence of a rigid cell wall located outside the plasmalemma. Unlike animal cells, which lack any external coat, the plant cell is surrounded by a complex polymeric framework based on cellulose, as well as hemicelluloses, pectins, and often lignin (Evert, 2006). The cell wall not only determines the shape of the cell and provides mechanical support to the whole plant, but also serves as a barrier protecting the protoplast from osmotic rupture upon water uptake (Graham et al., 2014). Animal cells, in contrast, usually have only a glycocalyx and cannot withstand significant osmotic pressure.
The second key difference is the presence in plants of plastids — double‑membrane organelles absent in animals. Most important are chloroplasts, which contain chlorophyll and carry out photosynthesis — the process of converting light energy into chemical energy, making the plant autotrophic (Beck, 2010). Derivatives of plastids are chromoplasts, which colour flowers and fruits, and leucoplasts (e.g., amyloplasts), which store starch (Evert, 2006). Animal cells, being heterotrophs, obtain energy solely by oxidizing ready‑made organic substances in mitochondria and do not contain plastids.
The third difference is the presence of a large central vacuole. In mature plant cells, the vacuole bounded by the tonoplast occupies up to 90% of the cell volume and performs many functions: maintaining turgor pressure, storing reserve substances (sucrose, proteins), depositing secondary metabolites and end products of metabolism, including pigments (anthocyanins) and calcium oxalate crystals (Andreev, 2001; Yakovlev et al., 2001). In animal cells, vacuoles are generally small (e.g., digestive and contractile vacuoles in protists) and do not play such a significant role in maintaining shape and metabolism.
The fourth difference concerns the organization of the cytoskeleton and cell division. Plant cells generally lack centrioles — structures that in animal cells participate in the formation of microtubule organizing centers of the spindle apparatus (Mauseth, 2017). Spindle formation in plant cells occurs differently. Furthermore, cytokinesis (division of the cytoplasm) in plants does not occur by furrowing (as in animals) but by the formation of a phragmoplast and a cell plate — a new cell wall that grows centrifugally and separates the daughter cells (Evert, 2006; Graham et al., 2014).
The fifth important difference is the mode of communication between cells. Adjacent plant cells are connected to each other by plasmodesmata — cytoplasmic channels passing through the cell walls (Serebryakova et al., 2006). This network unites the protoplasts of cells into a single whole — the symplast — providing rapid transport of ions and small molecules. Animal cells, lacking cell walls, are connected differently (e.g., via gap junctions), and their cytoplasm does not normally form such an extensive continuous compartment.
Thus, all the listed differences — cell wall, plastids, large vacuole, absence of centrioles, type of cytokinesis, and presence of plasmodesmata — are systemic traits that evolved to provide autotrophic nutrition, osmoregulation, and fixed growth of plants (Beck, 2010).
2. Fundamental properties of the plant cell
The vital activity of the plant cell is determined by several unique properties that distinguish it from heterotrophic eukaryotic cells and enable its autotrophic, sessile lifestyle. These properties — totipotency, osmotic activity (turgor), ability to synthesize a cellulose wall, and photoautotrophy — evolved together and are closely interrelated (Beck, 2010).
Totipotency. Totipotency is the ability of a somatic plant cell to realize the full genetic potential inherent to its species and, under certain conditions, to regenerate a whole organism (Fehér, 2019). Unlike animal cells, many differentiated plant cells (e.g., parenchyma cells) retain the ability to return to a meristematic state through dedifferentiation, which underlies methods of in vitro culture and vegetative propagation (Fehér, 2019). Totipotency is not a constitutive property of any somatic cell but rather an inducible state that requires reprogramming of gene expression and changes in the epigenetic landscape (Fehér, 2019). Detailed mechanisms of this phenomenon will be discussed in a separate article of the series.
Osmotic activity and turgor. The plant cell is an osmotic system in which the large central vacuole, filled with cell sap, is separated from the cytosol by the semipermeable tonoplast. Active accumulation of ions (K+, Na\+, Ca2+) and low‑molecular‑weight organic compounds (sucrose, proline) in the vacuole creates a more negative water potential compared to the environment, causing water uptake and the development of hydrostatic pressure against the cell wall — turgor (Andreev, 2001; Graham et al., 2014). Turgor provides tissue rigidity, maintains the shape of non‑lignified organs (leaves, stems), and is the driving force for cell expansion during growth (Evert, 2006). Regulation of turgor includes both passive water transport through aquaporins and active transport of ions and metabolites (Andreev, 2001). When dehydrated, cells lose turgor, which clinically manifests as plant wilting.
Ability to synthesize a cellulose cell wall. Unlike animal cells surrounded only by the plasmalemma, the plant cell forms an extracellular matrix — the cell wall — whose main structural component is cellulose (β‑1,4‑glucan) (Evert, 2006). Cellulose microfibrils are synthesized by enzyme complexes (cellulose synthase rosettes) located in the plasmalemma and are oriented by microtubules of the cytoskeleton (Mauseth, 2017). In addition to cellulose, the wall contains hemicelluloses, pectins, and (secondarily) lignin, which give it mechanical strength, elasticity, and protective properties. The presence of the wall not only determines cell shape but also restricts its motility, a key factor in the sessile lifestyle of plants (Graham et al., 2014).
Autotrophy. The plant cell is capable of photosynthesis — the process of converting sunlight energy into chemical energy of organic compounds (Beck, 2010). This ability is provided by specialized organelles — chloroplasts, containing chlorophyll a and b, as well as carotenoids, which are localized in the thylakoid membranes (Evert, 2006). During the light reactions, ATP and NADPH are synthesized, and in the chloroplast stroma, in the Calvin cycle, CO2 is fixed to form carbohydrates (Mauseth, 2017). Autotrophy makes the plant a primary producer of organic matter in ecosystems and determines its independence from external sources of organic nutrition.
Taken together, the listed fundamental properties — totipotency, turgor, ability to synthesize a cellulose wall, and autotrophy — are inextricably linked. For example, the existence of turgor as hydrostatic pressure is possible only in the presence of a rigid cell wall that counteracts the osmotic expansion of the protoplast. In turn, the cell wall, synthesized from products of photosynthesis, creates mechanical support for photosynthetic organs, and totipotency provides the regenerative potential necessary to replace damaged tissues during growth and development (Lee and Torii, 2026).
3. Structure and compartmentation of the plant cell

Structure of a plant cell
The living contents of a plant cell — the protoplast — are separated from the external environment by the cell wall and are characterized by complex internal organization. The protoplast is subdivided into functionally distinct compartments, allowing many biochemical processes to occur simultaneously and independently (Evert, 2006). The most important components of the protoplast are the plasmalemma, nucleus, cytoplasm (cytosol and organelles), vacuole, and the system of internal membranes.
Components of the plant cell are divided into several categories depending on their structural organization (Evert, 2006; Graham et al., 2014):
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Membrane‑bound organelles (common to eukaryotes): nucleus (bounded by a double membrane), endoplasmic reticulum (ER), Golgi apparatus (dictyosomes), mitochondria, peroxisomes, lysosome‑like structures (provacuoles).
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Membrane‑bound organelles (specific to plants): plastids (including chloroplasts, leucoplasts, chromoplasts) and the vacuolar system (tonoplast + central vacuole).
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Non‑membrane structures: ribosomes (free or bound to ER), microtubules, microfilaments (cytoskeleton), as well as inclusions such as starch grains, aleurone grains, calcium oxalate crystals, lipid droplets.
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Supramembranous structures: cell wall (primary and secondary), including cellulose microfibrils, hemicelluloses, pectins, and (upon lignification) lignin.
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Intercellular contacts: plasmodesmata — cytoplasmic channels piercing the cell walls and connecting the protoplasts of adjacent cells.
3.1. The protoplast and its boundaries
Protoplast (from Greek protos — first and plastos — molded, formed) is all the living contents of a plant cell, including the nucleus, cytoplasm, and organelles, but excluding the cell wall (Evert, 2006; Yakovlev et al., 2001). The protoplast is a complex colloidal system, the basis of which is water (60–90%), proteins, nucleic acids, lipids, and carbohydrates. All processes of metabolism, synthesis and breakdown of substances, as well as storage and transmission of hereditary information are localized in the protoplast.
The protoplast is bounded by two main membranes:
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Plasmalemma (plasma membrane, or ectoplast) — the outer membrane of the protoplast, directly adjacent to the cell wall. It is a universal membrane about 8–10 nm thick, built according to the “fluid mosaic” model: a lipid bilayer (mainly phospholipids and sterols) with integral and peripheral proteins embedded in it (Evert, 2006; Graham et al., 2014). Functions of the plasmalemma include:
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selective transport of substances into and out of the cell (via channels, carriers, and pumps, including H\+-ATPase, which creates a proton gradient);
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reception of signals (hormones, stress factors);
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participation in the synthesis and orientation of cellulose microfibrils (through integrated cellulose synthase complexes) (Mauseth, 2017).
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Tonoplast (vacuolar membrane) — the membrane that bounds the central vacuole and separates its contents (cell sap) from the cytosol. The tonoplast is similar in structure to the plasmalemma but differs in its set of transport proteins (aquaporins, H+-ATPase, H\+-pyrophosphatase, Na+/H\+ and Ca2+/H\+ antiporters, and metabolite transporters) (Andreev, 2001). Functions of the tonoplast: regulation of cytosolic ion homeostasis, maintenance of turgor, storage of secondary metabolites and waste products.
Between the plasmalemma and the tonoplast lies the cytosol (hyaloplasm) — a colloidal solution in which organelles and the cytoskeleton are suspended. The cytosol serves as the medium for glycolysis, fatty acid synthesis, part of nitrogen metabolism, and many other processes (Evert, 2006).
3.2. The nucleus
Main page: Plant cell nucleus
Nucleus (nucleus) — the central compartment of the eukaryotic cell, containing the vast majority of the genetic material and performing the functions of storing, reproducing, and realizing hereditary information (Yakovlev et al., 2001; Beck, 2010). A typical plant cell has one nucleus (rarely several); in highly specialized phloem sieve elements the nucleus degenerates at maturity (Evert, 2006).
The nucleus consists of the following structural elements:
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Nuclear envelope (karyolemma) — a double membrane (outer and inner) perforated by nuclear pores (nuclear pore complexes). The outer membrane is continuous with the endoplasmic reticulum and bears ribosomes. Pore complexes carry out selective transport of macromolecules (mRNA, ribosomal subunits, regulatory proteins) between the nucleoplasm and the cytosol (Evert, 2006).
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Nucleoplasm (karyolymph) — the semi‑fluid matrix of the nucleus, containing chromatin, nucleoli, and many enzymes (DNA polymerases, RNA polymerases, replication factors).
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Chromatin — a complex of linear double‑stranded DNA with nuclear proteins, primarily histones. In the interphase nucleus, more condensed heterochromatin (mostly inactive) and less condensed euchromatin (transcriptionally active) can be distinguished (Beck, 2010). Before cell division, chromatin condenses to form chromosomes visible under a light microscope.
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Nucleolus — a dense body without a membrane; the site of rRNA synthesis and assembly of ribosome precursors (small and large subunits). The number of nucleoli usually corresponds to the number of nucleolar organizers (secondary constriction regions of chromosomes).
Main functions of the nucleus:
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Storage of genetic information — the DNA in the chromosomes contains genes encoding all proteins and regulatory RNAs of the cell (Evert, 2006).
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Realization of genetic information — transcription of DNA occurs in the nucleus, producing various types of RNA (messenger, ribosomal, transfer, small nuclear, and small nucleolar RNAs) (Beck, 2010).
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DNA replication — duplication of genetic material in the S‑phase of the cell cycle, necessary to transfer identical copies of the genome to daughter cells.
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Regulation of the cell cycle and differentiation — the nucleus controls transitions between cell cycle phases (G₁/S and G₂/M) through the expression and activation of cyclin‑dependent kinases (CDKs) and their inhibitors (Sablowski and Gutierrez, 2022; Dewitte and Murray, 2003).
Thus, the nucleus acts as the command center of the cell, coordinating through signaling pathways (e.g., the retinoblastoma pathway, E2F/DP factors) the activity of all other compartments and determining the differentiation path of the cell (Lee and Torii, 2026).
3.3. The cytoplasm and its organization
Cytoplasm — the part of the protoplast enclosed between the plasmalemma, tonoplast, and nuclear envelope. It is a highly organized colloidal system in which the main processes of metabolism (protein synthesis, lipid synthesis, poly‑ and oligosaccharide synthesis, glycolysis, etc.) take place (Evert, 2006; Yakovlev et al., 2001). The cytoplasm is not a homogeneous mass but includes cytosol (the ground substance, or hyaloplasm) and numerous structures suspended in it — organelles and the cytoskeleton (Graham et al., 2014).
Cytosol and its properties
Cytosol (hyaloplasm) — the semi‑fluid, optically transparent phase of the cytoplasm, representing a colloidal solution of proteins, enzymes, metabolites, ions, and RNA. The cytosol contains up to 20–25% of all cellular proteins, including enzymes of glycolysis, the pentose phosphate pathway, fatty acid synthesis, and many other pathways (Evert, 2006).
Main functions of the cytosol: 1) providing a medium for metabolic reactions; 2) transport of metabolites between organelles by diffusion or flow (cyclosis); 3) maintenance of pH and ionic composition (together with the vacuole). The viscosity and fluidity of the cytosol can change depending on the physiological state of the cell.
Cytoskeleton
Main page: Plasma membrane and cytoskeleton
The cytoskeleton is a dynamic network of protein filaments that penetrate the cytosol. It provides mechanical support, determines cell shape, participates in intracellular transport and movement of organelles, and plays a key role in cell division (Mauseth, 2017; Beck, 2010). Two main types of filaments are distinguished:
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Microtubules — hollow cylindrical structures about 25 nm in diameter, built from heterodimers of α‑ and β‑tubulin. Microtubules have polarity and are capable of rapid polymerization/depolymerization (dynamic instability). In interphase cells, microtubules form a cortical array that directs the movement of cellulose synthase complexes in the plasmalemma and thereby determines the orientation of cellulose microfibrils (Mauseth, 2017). During mitosis, microtubules form the spindle apparatus, and during cytokinesis, the phragmoplast.
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Microfilaments (actin filaments) — thin (5–7 nm diameter) double helices of globular actin (G‑actin) polymerizing into filaments (F‑actin). Actin filaments are localized predominantly in the cortical layer of the cytoplasm and in transvacuolar strands. They participate in cytoplasmic streaming (cyclosis), mediated by the interaction of actin with myosin, as well as in the formation of the preprophase band (a precursor of the division zone) (Sablowski and Gutierrez, 2022; Graham et al., 2014).
Ribosomes
Ribosomes — non‑membrane organelles 15–25 nm in size, consisting of ribosomal RNA (rRNA) and proteins. Each ribosome is composed of two subunits (large and small). In the cytosol, ribosomes can be free or form polyribosomes (complexes of several ribosomes on one mRNA molecule) (Evert, 2006). Free ribosomes synthesize proteins destined for the cytosol, nucleus, mitochondria, peroxisomes, as well as ribosomal proteins (Graham et al., 2014). Ribosomes attached to the outer surface of the endoplasmic reticulum synthesize proteins that enter the ER lumen and are then directed to the Golgi apparatus, vacuoles, plasmalemma, or the extracellular space (Mauseth, 2017).
Endoplasmic reticulum (ER)
Main page: Endoplasmic reticulum and Golgi apparatus
Endoplasmic reticulum (ER) — a system of membranous cisternae, tubules, and vesicles connected to the outer membrane of the nuclear envelope. Two types of ER are distinguished (Evert, 2006; Beck, 2010):
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Granular (rough) endoplasmic reticulum (RER) — cisternae with ribosomes attached to their cytosolic surface. The RER is the site of synthesis of secretory, membrane, and vacuolar proteins, as well as their initial glycosylation (attachment of oligosaccharide chains). Protein folding and disulfide bond formation also occur in the RER lumen.
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Smooth (agranular) endoplasmic reticulum (SER) — lacks ribosomes and is developed in cells actively synthesizing lipids (including phospholipids, sterols, waxes, cutin, suberin) and carbohydrates. The SER participates in detoxification, accumulation of Ca2+ ions, and synthesis of some secondary metabolites (Mauseth, 2017).
Golgi apparatus (dictyosomes)
Golgi apparatus in plant cells is represented by individual stacks of flattened membranous cisternae called dictyosomes. A cell usually contains from a few to several tens of dictyosomes (Evert, 2006). Each dictyosome consists of 3–12 cisternae curved so that cis- (forming) and trans- (secretory) sides are formed. Functions of the Golgi apparatus (Evert, 2006; Graham et al., 2014):
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Modification, sorting, and packaging of proteins arriving from the RER (cleavage of signal peptides, phosphorylation of oligosaccharides to tag vacuolar proteins).
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Synthesis of cell wall matrix polysaccharides — pectins and hemicelluloses (e.g., xyloglucan). Polysaccharides are packaged into secretory vesicles.
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Formation of the cell plate during cytokinesis — Golgi vesicles carrying pectins and polysaccharides fuse at the equatorial plane, forming the middle lamella and new patches of plasmalemma.
Other cytoplasmic organelles
In addition to those listed, the cytoplasm of a plant cell contains:
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Mitochondria — double‑membrane organelles that carry out cellular respiration and ATP synthesis (more details in a separate article).
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Plastids (including proplastids, chloroplasts, leucoplasts, chromoplasts) — autonomous organelles with their own DNA, involved in photosynthesis, synthesis of starch, lipids, amino acids, and pigments (separate article).
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Peroxisomes (microbodies) — single‑membrane organelles containing catalase and oxidases. They participate in photorespiration (glycolate pathway) in leaves and in the conversion of fats to carbohydrates (glyoxysomes) in germinating seeds (Evert, 2006; Andreev, 2001).
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Spherosomes (lipid droplets) — small bodies surrounded by a half‑membrane, serving to store reserve lipids.
The entire cytoplasm is penetrated by a system of membranes and filaments, ensuring spatial and functional compartmentation, without which the simultaneous occurrence of diverse biochemical reactions would be impossible. Further description of individual organelles follows.
3.4. The cell wall
Cell wall — a rigid but elastic extracellular structure surrounding the protoplast and separated from it by the plasmalemma. It is a product of the vital activity of the protoplast and performs supportive, protective, transport, and signaling functions (Evert, 2006; Serebryakova et al., 2006). The cell wall is one of the key features distinguishing the plant cell from the animal cell; it provides mechanical strength to tissues, counteracts the osmotic pressure of the vacuole (turgor), and participates in intercellular interactions (Graham et al., 2014).
Chemical composition and structure. The basis of the cell wall is cellulose — a linear polysaccharide of β‑D‑glucose residues linked by β‑1,4‑glycosidic bonds. Cellulose molecules combine into microfibrils 10–25 nm in diameter, which are embedded in a matrix consisting of hemicelluloses (xyloglucan, xylan, mannan), pectins (pectates, pectinates), and glycoproteins (Mauseth, 2017). Pectins give the wall gel‑like properties and glue adjacent cells together, while hemicelluloses bind cellulose microfibrils to each other and to the pectin matrix (Evert, 2006). Secondary walls (in some cell types) are additionally impregnated with lignin — an aromatic polymer that gives the wall rigidity, hydrophobicity, and resistance to microbial degradation (Graham et al., 2014).
Two types of cell walls are distinguished (Yakovlev et al., 2001; Beck, 2010):
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Primary cell wall — formed during cell growth. It is thin (0.1–0.5 µm), contains up to 30% cellulose and many pectins, providing plasticity and the ability to stretch under turgor pressure. The primary wall is present in all living plant cells and is perforated by plasmodesmata.
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Secondary cell wall — deposited internally (between the plasmalemma and the primary wall) after cell growth ceases. It contains up to 40–60% cellulose, is often lignified, and may have a layered structure (S1, S2, S3). The secondary wall is characteristic of sclerenchyma fibers, xylem water‑conducting elements (tracheids and vessel elements), and stone cells (sclereids).
Pits and plasmodesmata. In the secondary wall, unthickened regions — pits — remain, through which the primary walls of adjacent cells remain in contact. A pair of two contiguous pits forms a pit field. In the pit region and in the primary wall, plasmodesmata pass — cytoplasmic channels connecting the protoplasts of adjacent cells (Evert, 2006). A plasmodesma is lined by the plasmalemma and contains a central rod (desmotubule) connected to the endoplasmic reticulum. Through plasmodesmata, intercellular transport of ions, small molecules (sugars, amino acids), as well as some proteins and RNAs occurs, making the tissues a single living system — the symplast (Graham et al., 2014; Sablowski and Gutierrez, 2022).
Thus, the cell wall is not an inert bag, but a dynamic, metabolically active structure, the synthesis of which is regulated by the cell and which is continuously modified in response to external and internal signals.
3.5. Interconnection of compartments
The plant cell functions as a single system due to a complex network of flows of substances, energy, and information between compartments. Each compartment (nucleus, cytosol, organelles, vacuole, cell wall) has its own set of enzymes and environmental conditions (pH, redox potential, ionic composition), which ensures compartmentation of metabolic pathways (Evert, 2006; Beck, 2010).
Vesicular transport. The main mechanism for directional movement of macromolecules and membranes between compartments is vesicular transport. Transport vesicles (coated with clathrin, COPI or COPII) bud from the endoplasmic reticulum and deliver proteins and lipids to the Golgi apparatus. In Golgi dictyosomes, the contents are modified, sorted, and packaged into secretory vesicles. These vesicles then fuse with the plasmalemma (exocytosis), releasing products (polysaccharides, glycoproteins) into the cell wall, or deliver hydrolases to vacuoles (Evert, 2006). The reverse process — endocytosis — allows the cell to take up extracellular material and recycle membrane components (Graham et al., 2014).
Cytoskeleton as a transport network. Microtubules and actin microfilaments serve as tracks for motor proteins (kinesins, dyneins, and myosins) that move vesicles and organelles (mitochondria, peroxisomes) to specific regions of the cell (Mauseth, 2017). For example, kinesins transport vesicles to the plus end of microtubules (cell periphery), while dyneins move them toward the minus end (centrosome — although plants lack centrioles, there are polar regions). The actin‑myosin system is responsible for cytokinesis and cytoplasmic streaming (cyclosis), which evenly distributes metabolites and organelles (Sablowski and Gutierrez, 2022).
Direct intercompartment exchange. Some metabolites and ions can move between compartments through specialized membrane carriers and channels without a vesicular step. For example, pyruvate and malate are transported across mitochondrial and chloroplast membranes; sugars and ions cross the plasmalemma and tonoplast via H\+-symporters and antiporters (Andreev, 2001). ATP synthesized in mitochondria and chloroplasts exits into the cytosol through pores or antiporters and supplies energy to all energy‑dependent processes.
Signaling pathways and regulation. Intercompartment interaction also occurs through intracellular signaling cascades. For example, an increase in cytosolic Ca2+ concentration (in response to a hormone or stress) can be initiated by Ca2+ release from the vacuole (via IP3-gated channels) or from the endoplasmic reticulum (Evert, 2006). The nucleus receives information about the state of the cell through retrograde signaling (e.g., from plastids — chloroplast biogenesis under the control of nuclear genes, as well as reverse signaling from organelles to the nucleus). These signals regulate gene expression, protein activity, and the cell cycle (Lee and Torii, 2026).
Thus, the functional unity of the plant cell is achieved through a dynamic balance of direct flows, vesicular traffic, cytoskeletal transport, and signaling networks coordinating the work of all compartments. Disruption of any of these links (e.g., blockade of vesicular transport due to mutations in SNARE proteins) leads to severe cytological and physiological defects (Sablowski and Gutierrez, 2022).
4. Types of plant cells (classification)
Cells of higher plants, despite a common structural plan, exhibit significant morphological and functional diversity, which is the result of differentiation during ontogeny. Depending on the functional state and tissue affiliation, several main types of cells are distinguished (Evert, 2006; Serebryakova et al., 2006).
4.1. Classification by functional state
Based on the ability to divide and the degree of specialization, two large groups are distinguished:
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Meristematic (undifferentiated) cells — cells of meristems (formative tissues), characterized by an isodiametric shape, dense cytoplasm, a large nucleus, small vacuoles, and a high level of mitotic activity (Evert, 2006). They retain the ability for unlimited division and give rise to all specialized cells of the plant. Meristems are localized in the shoot and root apices, in the cambium, and in the phellogen (Graham et al., 2014).
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Permanent (differentiated) cells — cells that have lost the ability to divide and acquired specific structures that enable them to perform particular functions (support, conduction, photosynthesis, storage, secretion, etc.) (Mauseth, 2017). Differentiation is accompanied by an increase in size, the appearance of a large vacuole, and changes in shape and cell wall properties.
4.2. Classification by tissue affiliation (functional)
In the plant body, several main types of cells are distinguished, which are grouped into the corresponding tissues (Yakovlev et al., 2001; Beck, 2010).
Parenchyma cells. The most widespread type, forming the bulk of the plant body. These are living, usually isodiametric cells with a thin primary wall. Parenchyma cells perform various functions:
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Assimilation parenchyma (chlorenchyma) — contains chloroplasts and serves as the main site of photosynthesis (leaf mesophyll).
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Storage parenchyma — accumulates starch, lipids, proteins, water (in tubers, root crops, seeds, fleshy fruit pulp).
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Conducting parenchyma — participates in radial transport of substances (e.g., cells of medullary rays in wood).
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Aerenchyma (air parenchyma) — has large intercellular spaces, providing ventilation to submerged and wetland plants.
Collenchyma cells. Living, elongated cells with unevenly thickened primary walls (thickenings located at the corners or along tangential walls). They provide mechanical strength to young, growing organs (stems, petioles, leaf veins) and are capable of stretching together with surrounding tissues (Evert, 2006). Found mainly in dicotyledonous plants.
Sclerenchyma cells. Dead (protoplast‑lost) cells with evenly thickened, often lignified secondary walls. They are subdivided into:
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Fibers — long, narrow, tapered prosenchyma cells. They form strong strands (bast fibers, wood fibers) and serve a supporting function (Graham et al., 2014).
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Sclereids (stone cells) — short, isodiametric or irregularly shaped cells with very thick walls (e.g., in seed coats, nut shells, pear fruit flesh) (Mauseth, 2017).
Conducting cells. Specialized for transport of water and minerals (xylem) or organic assimilates (phloem).
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Xylem (wood): water‑conducting elements — tracheids (dead prosenchyma cells with bordered pits, characteristic of gymnosperms and ferns) and vessel elements (shorter and wider, with perforation plates on the end walls, characteristic of angiosperms) (Evert, 2006).
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Phloem (bast): sieve elements — living but enucleate cells. In angiosperms — sieve tube elements with sieve plates, accompanied by companion cells; in gymnosperms — sieve cells (Mauseth, 2017).
Secretory and excretory cells. Cells that produce and/or accumulate secondary metabolites (essential oils, resins, gums, alkaloids, nectar) or remove waste products. They can be single idioblasts (e.g., cells with calcium oxalate crystals, essential oil cells) or be organized into structures (laticifers, resin ducts, nectaries, hydathodes) (Evert, 2006; Graham et al., 2014).

Paths of differentiation and regeneration of plant cells
Generalized scheme of terminology used to describe changes in cell fate in plants: differentiation, dedifferentiation (broad and narrow sense), transdifferentiation, as well as pathways of callus formation and shoot/root regeneration. Fehér, 2019, figure 1, CC BY 4.0 <a class="common-share-detail" rel='nofollow' href='https://creativecommons.org/licenses/by/4.0/' aria-label='Common Share CC BY 4.0' target='_blank'>CC BY 4.0</a>.
The above classification reflects the final states of differentiated cells. However, in plant ontogeny, all specialized cells originate from meristematic cells. The process of differentiation is accompanied not only by the loss of the ability to divide and the acquisition of a specific structure but also, in most cases, by the loss of totipotency — the ability to regenerate a whole organism. Nevertheless, under certain conditions (e.g., upon wounding or in in vitro culture), differentiated cells can undergo dedifferentiation, returning to a meristematic state, or even transdifferentiation, changing one type of specialization for another (Fehér, 2019). The ability to regenerate a whole plant from a somatic cell underlies the phenomenon of totipotency, which is widely used in plant biotechnology (micropropagation, production of virus‑free material, somatic embryogenesis). The mechanisms of differentiation, maintenance, and reprogramming of the cellular state, including epigenetic regulation and the role of the retinoblastoma pathway, will be discussed in detail in a separate article of the “Fundamentals of cytodifferentiation and totipotency” block (Lee and Torii, 2026; Sablowski and Gutierrez, 2022).
5. Cell cycle and division
Main page: Cell division
The life cycle of a dividing somatic cell encompasses two sequential processes: interphase (preparation for division) and mitosis (nuclear division), usually followed by cytokinesis (division of the cytoplasm and formation of two daughter cells) (Dewitte and Murray, 2003; Francis, 2007). Unlike animal cells, plants lack centrioles, and cytokinesis occurs by forming a cell plate rather than by furrowing (Evert, 2006).
5.1. Interphase
Interphase typically occupies at least 90% of the cell cycle time and includes three periods (Sablowski and Gutierrez, 2022):
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G₁ phase (postmitotic) — the cell grows; RNA, proteins (including enzymes for DNA replication) are synthesized; the number of organelles increases. At the end of G₁, a checkpoint determines readiness for DNA replication.
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S phase (synthetic) — DNA replication occurs, as a result of which each chromosome becomes composed of two sister chromatids, and the ploidy of the nucleus increases from 2C to 4C (Francis, 2007). Simultaneously, histones and proteins required for replication are synthesized.
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G₂ phase (premitotic) — synthesis of proteins and RNA continues; components necessary for mitosis (including tubulins) accumulate. At the end of G₂ is a second checkpoint, verifying the completeness and accuracy of DNA replication.
5.2. Mitosis (karyokinesis)
Mitosis is the process of distributing replicated chromosomes between two daughter nuclei. In higher plants, four phases are distinguished (Evert, 2006; Mauseth, 2017):
Prophase. Chromatin condenses; chromosomes become visible under a light microscope (each consisting of two sister chromatids joined at the centromere). The nucleolus disappears, and the nuclear envelope fragments and is incorporated into the endoplasmic reticulum. In the cytoplasm, a preprophase band of microtubules and actin filaments forms, girdling the nucleus in the zone of the future cell plate (Sablowski and Gutierrez, 2022).
Metaphase. The nuclear envelope has completely disappeared. Chromosomes align at the equatorial plane of the spindle apparatus formed by microtubules. Spindle fibers attach to the kinetochores of the centromeres.
Anaphase. Sister chromatids separate and, with the help of the spindle apparatus, move to opposite poles of the cell. Each chromatid becomes a daughter chromosome.
Telophase. At each pole, a new nuclear envelope forms; chromosomes decondense; nucleoli reappear. Spindle microtubules depolymerize.
5.3. Cytokinesis in plants
Cytokinesis begins in telophase and has features that distinguish it from cytokinesis in animals (Evert, 2006; Graham et al., 2014):
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In the equatorial region (between the two daughter nuclei), microtubules and actin filaments form the phragmoplast — a cylindrical structure that guides the movement of Golgi‑derived vesicles.
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Vesicles containing pectins and hemicelluloses fuse to form the cell plate — a new patch of cell wall that grows centrifugally (from the center toward the periphery).
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The cell plate reaches the mother cell wall and fuses with it, dividing the protoplast into two daughter cells. Plasmodesmata — channels connecting the cytoplasm of adjacent cells — form in the plate and adjacent new walls.
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Cytokinesis results in two cells, each with an identical set of chromosomes (2n) and a volume of cytoplasm that is then restored during the G₁ phase.
5.4. Reference to the article on division
Detailed mechanisms of cell cycle regulation (cyclin‑dependent kinases, cyclin‑dependent inhibitors, retinoblastoma pathway) and specifics of mitosis in various plant tissues will be discussed in a separate article of the “Plant cell division” block. It is important to emphasize here that mitotic division underlies growth, regeneration, and vegetative propagation, and its correct progression is controlled by a complex signaling network (Lee and Torii, 2026; Sablowski and Gutierrez, 2022).
6. Plant cytology as a science
Cytology (from Greek kytos — cell and logos — study) is a branch of biology that studies the structure, chemical composition, functions, development, and reproduction of cells. In its application to plants, cytology is a fundamental basis for understanding tissue organization, physiological processes, and biotechnological approaches in agronomy (Evert, 2006; Yakovlev et al., 2001).
Object and subject of cytology. The object of cytology is the plant cell at all levels of structural organization — from the intact protoplast to the molecular organization of membranes and the cytoskeleton. The subject of research includes: 1) morphology and ultrastructure of cellular compartments; 2) physicochemical properties of the cytoplasm and its organelles; 3) the cell cycle, mitosis, and meiosis; 4) differentiation, totipotency, and cellular aging; 5) cell responses to external signals and stress (Mauseth, 2017; Graham et al., 2014).
Historical outline. The foundations of cytology were laid in the second half of the 17th century, when R. Hooke (1665) first described cells in a slice of cork. In 1831, R. Brown discovered the nucleus, and in 1838–1839, M. Schleiden and T. Schwann formulated the cell theory, asserting that all living organisms are composed of cells (Serebryakova et al., 2006). Later, R. Virchow (1855) added the principle “omnis cellula e cellula” — every cell arises from another cell. In the second half of the 19th — early 20th century, mitochondria (R. Altmann, 1890; K. Benda, 1898), plastids (A. Schimper, 1883), the Golgi apparatus (C. Golgi, 1898) were discovered, and mitosis was described (E. Strasburger, W. Flemming). A landmark achievement was the discovery of the DNA structure (J. Watson and F. Crick, 1953), which enabled the transition to molecular cytology (Beck, 2010).
Main methods of cytology. Progress in cell science is inextricably linked to the development of microscopic and biochemical methods (Evert, 2006; Graham et al., 2014):
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Light microscopy — allows observation of living and fixed cells at magnifications up to 1500×, revealing the nucleus, plastids, vacuoles, and chromosomes (after staining). Variants include phase‑contrast, fluorescence (including the use of fluorescent proteins such as GFP), and confocal laser scanning microscopy.
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Electron microscopy — provides resolution down to 0.2 nm, allowing visualization of the ultrastructure of membranes, ribosomes, microtubules, and organelle details. Transmission electron microscopy (TEM) is used to study the internal structure of sections, while scanning electron microscopy (SEM) is used to examine the surface of cells and tissues.
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Cytochemistry and immunocytochemistry — specific staining or antibody labeling allows localization of particular substances (DNA, RNA, proteins, polysaccharides, ions) within the cell. Fluorescent protein tagging (GFP and its derivatives) and in situ hybridization methods (FISH) enable visualization of gene expression and chromosome positions.
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Methods of cell biology — density‑gradient centrifugation for organelle isolation, culture of isolated protoplasts, microinjection, electrophysiology (patch‑clamp) for studying ion channels.
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Molecular cytological approaches — flow cytometry for DNA content analysis, chromosome microdissection, genome editing (CRISPR/Cas) for analyzing gene functions in the cell.
Significance for agricultural sciences. Cytological research underpins plant breeding (karyotype analysis, identification of polyploid forms, cytogenetic control of hybrids), biotechnology (clonal micropropagation, somatic hybridization, production of haploid plants), phytopathology (diagnosis of viral and bacterial infections by intracellular inclusions, studying pathogen–plant cell interactions), and stress physiology (cytological markers of drought and salt tolerance, assessment of membrane damage) (Mauseth, 2017; Fehér, 2019).
Thus, plant cytology is a comprehensive discipline that combines structural, functional, genetic, and molecular approaches to the study of the cell. Its achievements create a theoretical basis for increasing the productivity of agricultural crops, plant resistance to stress, and the implementation of modern biotechnologies.
7. Current state and prospects of plant cytology
Modern plant cytology is undergoing a stage of integration of classical microscopic methods with advanced technologies of molecular biology, biophysics, and bioinformatics. This allows cellular processes to be studied not only statically but also dynamically — in living cells and tissues, and with unprecedented spatiotemporal resolution (Evert, 2006; Graham et al., 2014).
New methods and approaches. The key technological drivers of modern cytology are:
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Confocal laser scanning microscopy (CLSM) and super‑resolution microscopy (STED, PALM, STORM) — allow visualization of structures down to 20–50 nm, bypassing the diffraction limit of the light microscope. These methods are used to study cytoskeleton dynamics, vesicle movement, and the spatial organization of receptors in the plasmalemma (Evert, 2006; Lee and Torii, 2026).
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Cryo‑electron tomography (cryo‑ET) — enables three‑dimensional reconstructions of the ultrastructure of cells in their native, frozen state, without chemical fixation. This method is particularly important for studying the fine structure of synapses, plasmodesmata, the Golgi apparatus, and chloroplasts (Beck, 2010).
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Fluorescent protein tagging (GFP, RFP, mCherry, and photoactivatable proteins) combined with genetic engineering methods allows tracking the localization and dynamics of any protein in living cells. For example, visualization of microtubules with MAP4‑GFP or actin with Lifeact‑GFP has revealed fine mechanisms of cytokinesis and polar growth (Sablowski and Gutierrez, 2022).
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Flow cytometry and fluorescence‑activated cell sorting (FACS) — provide rapid analysis and isolation of individual cells or nuclei based on their ploidy, size, and fluorescent markers. Combined with single‑nucleus RNA sequencing (snRNA‑seq), this approach allows mapping the transcriptomes of all cells of a root or leaf at sufficient resolution to reconstruct cellular trajectories (Lee and Torii, 2026; Sablowski and Gutierrez, 2022).
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Genome editing (CRISPR/Cas9) — opens the possibility of targeted modification of genes regulating the cell cycle, differentiation, and stress response. Plant lines with knockouts and knock‑ins of cyclin, CDK, and KRP genes have been created, allowing functional characterization of many division regulators (Fehér, 2019; Dewitte and Murray, 2003).
Current research directions. At the forefront of plant cytology are the following problems:
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Mechanisms of polarity and asymmetric division — how external signals (auxin, brassinosteroids, mechanical stress) and intracellular polarity proteins (BASL, BRXL2, SOSEKI) determine the division plane and unequal distribution of cell‑fate determinants (Lee and Torii, 2026; Sablowski and Gutierrez, 2022).
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Cytoskeleton and intracellular transport — deciphering the signaling pathways linking microtubules and actin to motor proteins, and elucidating the roles of these systems in gravitropism, photoperiodism, and immune responses (Mauseth, 2017).
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Endomembrane system and vesicular traffic — studying the formation and maturation of endosomes, multivesicular bodies, autophagosomes, and their roles in membrane recycling, protein degradation, and pathogen resistance (Andreev, 2001).
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Compartmentation and signaling — how the cell coordinates the work of the nucleus, chloroplasts, mitochondria, and peroxisomes through retrograde signaling and how these connections are disrupted under stress (Beck, 2010).
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Totipotency and reprogramming — identification of transcription factors and epigenetic modifications necessary for dedifferentiation and regeneration of a whole plant from a single somatic cell. The creation of single‑cell gene expression atlases has made it possible to discover rare transitional states (Fehér, 2019; Lee and Torii, 2026).
Significance for agricultural sciences. Modern cytology is directly implemented in agricultural practice:
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Breeding and seed production. Cytogenetic analysis (karyotyping, FISH, GISH) is used to control hybridization, identify polyploid and aneuploid forms, and detect chromosomal rearrangements, which is critically important in the development of new varieties of wheat, triticale, sugar beet, and sunflower (Yakovlev et al., 2001).
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Biotechnology. Tissue culture and isolated protoplast methods, based on totipotency, allow propagation of valuable genotypes (micropropagation), virus elimination (apical meristem culture), production of somatic hybrids (protoplast fusion), and haploid plants (embryo culture, anther culture) (Fehér, 2019).
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Phytopathology and stress tolerance. Cytological markers (membrane damage, chromatin aggregation, reactive oxygen species production) are used to screen varieties for resistance to pathogens, herbicides, salinity, and drought. Plasmodesmata and vesicle traffic are targets for viruses, and understanding these interactions helps create virus‑resistant plants (Andreev, 2001; Mauseth, 2017).
Open questions. Despite enormous progress, fundamental mysteries remain: how exactly does a cell assess its size and coordinate division with growth; what molecular switches determine the choice between mitosis and endoreduplication; how do plasmodesmata regulate their permeability, and how are signals transmitted from the cell wall to the nucleus? Answers to these questions will be obtained using new technologies — from super‑resolution microscopy to artificial intelligence for analyzing large image datasets (Sablowski and Gutierrez, 2022; Lee and Torii, 2026).
References
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Andreev, I.M. 2001. The role of the vacuole in the regulation of plant cell metabolism. Russian Journal of Plant Physiology, 48(5), pp. 672–683.
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Beck, C.B. 2010. An Introduction to Plant Structure and Development: Plant Anatomy for the Twenty‑First Century. 2nd ed. Cambridge: Cambridge University Press, pp. 3–57 (Chapter 3: The protoplast of the eukaryotic cell).
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Dewitte, W. and Murray, J.A.H. 2003. The plant cell cycle. Annual Review of Plant Biology, 54, pp. 235–264. DOI: 10.1146/annurev.arplant.54.031902.134836 PubMed
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Evert, R.F. 2006. Esau’s Plant Anatomy: Meristems, Cells, and Tissues of the Plant Body – Their Structure, Function, and Development. 3rd ed. Hoboken: John Wiley & Sons, pp. 15–41 (Chapter 2: The protoplast: plasma membrane, nucleus, and cytoplasmic organelles).
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Fehér, A. 2019. Callus, dedifferentiation, totipotency, somatic embryogenesis: what these terms mean in the era of molecular plant biology? Frontiers in Plant Science, 10, p. 536. DOI: 10.3389/fpls.2019.00536 PubMed
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Francis, D. 2007. The plant cell cycle – 15 years on. New Phytologist, 174(2), pp. 261–278. DOI: 10.1111/j.1469-8137.2007.02038.x PubMed
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Graham, L.E., Graham, J.M. and Wilcox, L.W. 2014. Plant Biology. 2nd ed. Boston: Pearson, pp. 47–74 (Chapter 4: Cells).
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Lee, L.R. and Torii, K.U. 2026. Cell cycle in plant development and reprogramming. Development, 153(3), dev205318. DOI: 10.1242/dev.205318 PubMed
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Mauseth, J.D. 2017. Botany: An Introduction to Plant Biology. 6th ed. Burlington: Jones & Bartlett Learning, pp. 4–74 (Chapter 3: Cell Structure).
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Sablowski, R. and Gutierrez, C. 2022. Cycling in a crowd: coordination of plant cell division, growth, and cell fate. The Plant Cell, 34(1), pp. 193–208. DOI: 10.1093/plcell/koab222 PubMed
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Serebryakova, T.I., Voronin, N.S., Elenevsky, A.G., Batygina, T.B., Shorina, N.I. and Savinykh, N.P. 2006. Botanika s osnovami fitotsenologii: Anatomiya i morfologiya rasteniy [Botany with fundamentals of phytocenology: Plant anatomy and morphology]. Moscow: Akademkniga, pp. 53–100 (Chapter 2: Fundamentals of plant anatomy). (In Russian)
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Yakovlev, G.P., Chelombitko, V.A. and Dorofeev, V.I. 2001. Botanika: Uchebnik dlya vuzov [Botany: Textbook for universities]. St. Petersburg: SpetsLit, pp. 5–32 (Chapter 1: Cytology). (In Russian)



