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5 Cellular Structure andFunction
Mitochondrion
Golgi
endoplasmic
Centrioles
52
Free
ribosomes
Lysosome
Endosome
Peroxisome
Cytoskeleton
Figure5.1 Cell structures and organelles (created in BioRender).
apparatus
Plasma
membrane
Nucleus
Nucleolus
Rough
endoplasmic
reticulum
Smooth
reticulum
Microtubules
backdrop for intracellular processes not confined within membrane- bound organelles, the cellular
cytoplasm significantly impacts essential cellular functions. These functions encompass protein
folding, enzymatic catalysis, intracellular signalling, molecular transport and the spatial arrangement of molecules and organelles(2).
The plasma membrane, serving as a fundamental perimeter organelle, plays a pivotal role as a
selective barrier, distinguishing and isolating cellular components from their surroundings. This
membrane constitutes a primary hallmark of cellular existence and consists of three principal
constituents: lipids, proteins and carbohydrates, with their respective mass ratios varying according to the specific cell type. The membrane’s architecture, the physicochemical attributes of its
macromolecular components and their interplay collectively govern vital functions, including
selective transport, cell recognition, signalling and the compartmentalisation of intracellular
processes(3, 4).
5.2.2 Nucleus
The nucleus, one of the cell’s largest organelles, orchestrates a plethora of critical cellular functions, housing distinct subdomains with unique structural configurations and biochemical compositions(5). Foremost among these subdomains is the nucleolus, responsible for ribosomal RNA
transcription, processing and preribosomal assembly. Additionally, the nucleolus plays a pivotal
role in genome organisation during interphase. It regulates the stability of p53, a crucial transcription factor involved in cell cycle control, especially in response to DNA damage(6). Surrounded by
the nuclear envelope, a bilayered structure comprising the outer and inner nuclear membrane, the
nucleus maintains its integrity within a nucleoplasmic environment separated from the cytoplasmic milieu(7–9). The nuclear lumen contains chromatin, a complex of genomic DNA entwined
with RNAs, histones and various chromatin proteins(10, 11).
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5.2 Cell Structure andFunction
5.2.3 Ribosomes
Ribosomes, large ribonucleoprotein complexes, play a central role in protein synthesis across all
living cells(12). Comprising two subunits of unequal size, these molecular machines orchestrate
the translation process. The smaller ribosomal subunit engages with messenger RNAs (mRNAs),
decoding the genetic message by selecting appropriate aminoacyl- transfer RNA (tRNA) molecules(13). Meanwhile, the larger subunit harbours the peptidyl transferase centre, a catalytic site
that facilitates the formation of peptide bonds, thereby stringing together amino acids delivered by
tRNAs into a growing polypeptide chain(14, 15).
Emerging polypeptide chains exit the ribosome via a tunnel within the larger subunit and
subsequently interact with various protein factors. These factors play essential roles in enzymatic
processing, targeting and the membrane insertion of nascent chains, all of which occur at the
ribosomal tunnel’s exit(14, 15).
5.2.4 Mitochondria
Mitochondria, recognised as the cellular powerhouses within eukaryotic cells, hold pivotal significance in adenosine triphosphate (ATP) synthesis via oxidative phosphorylation. They are indispensable components in cellular energy metabolism and various signalling pathways. Mitochondria
are intricately organised into four distinct compartments: the outer membrane, intermembrane
space, inner membrane and matrix. Notably, the inner mitochondrial membrane, with its significantly greater surface area in comparison to the outer membrane, forms cristae invaginations that
house the oxidative phosphorylation system(16–18). Mitochondria fulfil crucial roles in maintaining cell viability across diverse growth conditions. These functions encompass activities such as
iron–sulfur cluster synthesis, mitochondrial protein import and maturation(19) and the management of a comprehensive genetic system, which includes the mitochondrial genome, essential
genome maintenance and regulatory factors, along with unique mitochondrial ribosomes distinct
from their cytosolic counterparts(20).
53
5.2.5 Peroxisomes
Peroxisomes, intracellular organelles encased by a singular lipid bilayer membrane enfolding a
dense protein matrix, serve critical roles in diverse cellular processes(21). These functions encompass the β- and α- oxidation of fatty acids, decomposition of hydrogen peroxide and the synthesis of
ether- phospholipids and docosahexaenoic acids(22). Peroxisomes engage in intricate interactions
with various cellular organelles, including mitochondria, the endoplasmic reticulum (ER) and
lipid droplets (LDs)(21, 23).
5.2.6 Vesicles andVacuoles
Vesicles, vacuoles and membranous sacs serve as integral components for storage and transport
within cells. Vesicles naturally form during secretion (exocytosis), uptake (endocytosis) and
intracellular material transport at the plasma membrane(24). Extracellular vesicles, comprising
microvesicles, exosomes, apoptotic bodies and argosomes are categorised based onsize, content, synthesis and function. Microvesicles, ranging from 100 to 1000 nm withdiverse shapes,
result from the regulated release through outward budding of the plasma membrane. Exosomes,
smaller vesicles (40–100 nm), play a role in intercellular communication, irrespective of
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5 Cellular Structure andFunction
54
cellular distance. They are present in biological fluids such as plasma, serum and breast milk,
formed from multivesicular bodies through inward budding of the endosome membrane.
Apoptotic bodies, measuring 1000–5000 nm, represent the ultimate outcome of cellular
fragmentation, containing intact organelles, DNA and histones. Lastly, argosomes are exosomelike vesicles carrying morphogens, proteins forming tissue concentration gradients involved in
signal transduction, thereby indicating cellular position during multicellular organism
development(25).
LDs represent specialised cytosolic lipid accumulations present in all organisms(26). These LDs
comprise a neutral lipid core enveloped by a monolayer membrane composed of phospholipids
and housing specific functional proteins(27, 28). LDs play pivotal roles in numerous fundamental
cellular processes, both in states of health and pathology(29). These functions encompass cell
signalling(30, 31), the regulation of cell fate in mature muscle stem cells(32), membrane trafficking, the production of inflammatory mediators(33–35) and the management of cellular stress and
homeostasis(36, 37). Beyond these fundamental roles, LDs are intricately involved in broader
physiological and pathological cellular phenomena, including participation in neoplastic processes associated with various malignancies(38), the sequestration and potential metabolism of
lipophilic toxins(39), resistance to anticancer drugs and the capacity of cancer cells to endure
chemotherapeutic stress(40).
5.2.7 Centrosome
The centrosome, the principal microtubule- organising centre in animal cells, exerts regulatory
control over cell motility, adhesion and polarity during interphase. It also orchestrates spindle pole
organisation during mitosis(41). Comprising two centrioles ensconced within an electron- dense
matrix called the pericentriolar material, the centrosome is unique among organelles due to its
lack of a bounding membrane. This feature allows for dynamic changes throughout the cell cycle
and close interactions with cytoplasmic components, primarily mediated through microtubules
organised and reorganised by centrosomes during different cell cycle phases(42). Centrosomes are
pivotal in various cellular processes, such as mitotic spindle formation, cell cycle progression,
neurogenesis, cell polarity and cell migration. Dysregulation of centrosomal functions can give rise
to a range of diseases, including cancer(43).
5.2.8 Lysosomes
Lysosomes, enclosed within a membrane, serve as vital cytoplasmic organelles responsible for the
degradation of a diverse array of biological macromolecules, encompassing proteins, lipids, carbohydrates and nucleic acids(44). Traditionally regarded as the ‘garbage- disposal system’ of the cell
due to their terminal degradation function, lysosomes have demonstrated involvement in various
other cellular processes. These include roles in metabolic signalling, gene regulation, immunity,
plasma membrane repair and cell adhesion and migration. Lysosome number, composition and
functions exhibit adaptability in response to environmental cues, as well as cellular and organismal requirements. They engage in physical and functional interactions with other cellular structures, forming membrane contact sites and display dynamic attributes such as mobility, alterations
in size and shape and undergoing fusion or fission events. Dysregulation of lysosomal function has
been associated with the pathogenesis of common diseases, including neurodegenerative and metabolic disorders, along with cancer(45).
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5.2 Cell Structure andFunction
5.2.9 Endoplasmic Reticulum
The ER, occupying a substantial portion of the cytoplasm, serves as the primary cellular
sitefor lipid and carbohydrate conjugate biosynthesis. Additionally, it plays a crucial role in
the folding, assembly and biosynthetic transport of secreted proteins and integral membrane
proteins(46).
The rough ER, distinguished by the presence of ribosomes on its cytoplasmic surface, is the
region specialised for protein synthesis, folding and degradation. Both soluble and transmembrane proteins are dispatched from the ER at designated sites known as ER export domains.
These domains consist of tubulovesicular membranes lacking ribosomes and bud- off vesicle
intermediates for transport to the Golgi apparatus. In contrast, the smooth ER comprises
ribosome- free tubular elements and is primarily involved in enzyme pathways related to drug
metabolism, steroid synthesis and calcium uptake and release(6).
5.2.10 Golgi Apparatus
The Golgi stack, comprising 4–8 flattened membrane cisternae, is functionally polarised. The
cis face receives cargo from the ER, while the trans face serves as the exit site for cargo destined
for various cellular destinations (47). Individual Golgi stacks merge through lateral fusion,
forming a twisted ribbon structure typically located centrally or juxtanuclear in most cell
types (48). This positioning is maintained by the radial centrosomal microtubule array in
non- polarised cells. During interphase, the Golgi forms a ribbon, but this structure disassembles into vesicles and tubules during mitosis to facilitate the partitioning of Golgi membranes between daughter cells (49). The Golgi apparatus plays a crucial role in regulating
numerous high- level cellular functions, including cell polarisation, DNA repair, apoptosis,
stress responses, mitosis (50), directed migration, secretion, metabolism, autophagy and
inflammation(47).
55
5.2.11 Cytoskeleton
The cytoskeleton is a multifaceted, fibrous reticular structure with diverse functions(51). It encompasses several cytoskeletal components, including actin filaments, microtubules, intermediate
filaments and spectrin. Two main regions within the cell house the cytoskeleton: the cortical
cytoskeleton, located beneath the cell membrane and the cytoplasmic cytoskeleton. The cortical
cytoskeleton provides structural support, flexibility and elasticity to the cell membrane, contributing to changes in membrane shape under external forces. The cytoplasmic cytoskeleton plays
essential roles in intracellular transport, cell stabilisation and force transmission(52). Coordinated
interactions between cortical and cytoplasmic cytoskeletal proteins are pivotal in various cellular
processes, including the regulation of cell motility, morphological changes, pinocytosis, endocytosis, cell adhesion and signal transduction(53, 54).
Cytoskeletal proteins are distributed in the cytoplasm, and nucleoskeletal proteins are present in
the nucleus. The linker of nucleoskeleton and cytoskeleton complex, spanning the nuclear envelope, acts as a bridge connecting the cytoskeleton to the nucleoskeleton, facilitating signal communication between the nucleus and cytoplasm. Additionally, it plays a crucial role in maintaining
the nuclear envelope’s architectural integrity(55).
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5 Cellular Structure andFunction
Cell–cell junctions
Connected to cytoskeleton
intermediate filaments
intermediate filaments
Gap junctions
56
Adherence junctions
by actins
Anchoring junctions
Desmosomes Hemidesmosomes Focal adhesions
Connected to
cytoskeleton by
Connected to
cytoskeleton by
Connected to
cytoskeleton by actins
Tight junctions
Connected to cytoskeleton by
actins
Figure5.2 Types of cell junctions. Source: Modified from(56); created in BioRender.
5.3 Intercellular Junctions
Intercellular junctions are specialised regions of the plasma membrane characterised by unique
structural and biochemical features that enable specific interactions between neighbouring
cells(56). Originally named based on their morphology and presumed functions, these junctions
play a crucial role in maintaining a tight barrier to prevent pathogen invasion and systemic
spread(57). There are three main functional categories of cell junctions; anchoring, tight and gap
junctions (see Figure5.2)(58). Adherence junctions and desmosomes provide essential adhesive
and mechanical properties that contribute to barrier function (59). Tight junctions hold cells
together and create an almost impermeable intercellular seal by fusing adjacent cell membranes(60) and primarily regulate paracellular transport, often representing the rate- limiting step
in this process(59). Gap junctions enable the passage of small molecules between adjacent cells
through channel- like structure. In addition to these primary junctions, hemidesmosomes are
integrin- based adhesive junctions linking cytoplasmic intermediate filaments to the basal lamina.
Integrins associated with cytoplasmic actin filaments form focal adhesions connecting cells to the
ECM(56).
5.4 Cell Communication andSignalling Pathways
Cell- to- cell communication plays a pivotal role in processes such as cell differentiation, morphogenesis, cell growth and homeostasis in multicellular organisms. Several molecular mechanisms
underpin cell communication, including cytoplasmic bridges, exosomes, direct interactions
between membrane proteins on adjacent cells and soluble messenger molecules that influence
target cells, both nearby and distant.
Cytoplasmic bridges encompass various structures, including gap junctions (61), tunnelling
nanotubes(62) and those formed during incomplete cytokinesis(63). Exosomes and ectosomes are
distinct types of extracellular vesicles produced by various cell types(64). They carry cargoes of
proteins, lipids and nucleic acids that impact transcription factors, signalling proteins and enzymes
in target cells(65).
Cell–cell adhesion and communication are facilitated by membrane multidomain proteins(66),
which often adopt stable conformations(67). In animals, cell contact is also mediated through the
ECM, featuring proteins like collagens, fibronectins, laminins(68, 69) and oligosaccharides(70)
interacting with cell membrane proteins such as cadherins, integrins, cell adhesion molecules and
selectins(71, 72).
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5.6 The Cell Cycle
Most cells produce soluble messenger molecules with the ability to influence distant cells. These
mediators encompass hormones, neuromediators, cytokines, growth factors and morphogens.
Target cells can be in close proximity to the mediator- emitting cell or at a considerable distance
within another individual(73).
Mediator receptors bridge the gap between intercellular communication and intracellular signalling. They belong to two main families: membrane receptors with binding sites on the external
cell surface and intracellular receptors that act on DNA (nuclear receptors in eukaryotes). The first
group’s receptors bind to mediators that do not enter the cell, while the second group’s receptors
respond to ligands that can penetrate the cell. Mediator receptors can be soluble intracellular transcription factors or proteins integrated into the plasma membrane. The nuclear receptors form a
significant family of related transcription factors(74). Plasma membrane receptors include channel receptors allowing specific ion influx(75), receptors with intrinsic enzymatic activities(76) and
receptors that activate various downstream intracellular effectors(77–79).
5.5 Interactions withthe Extracellular Matrix
The ECM encompasses diverse macromolecules that create the microenvironment surrounding
cells, offering mechanical and structural support. The interaction between cells and the ECM is
crucial in cellular behaviours like migration, growth and differentiation. The effects of ECM
macromolecules on cell signalling occur directly or indirectly(80). In direct regulation, ECM macromolecules engage with receptors like integrins(81) and CD44 or syndecans(82, 83). In indirect
regulation, ECM macromolecules collaborate with several receptor molecules and growth factors(84). Additionally, proteolytic degradation of ECM macromolecules can release matrix- bound
growth factors, activating their signalling pathways (85–87). This reciprocal communication
between ECM macromolecules and cells underpins processes such as ontogenesis, wound healing
and tissue homeostasis, while disturbances in ECM structure, metabolism or ECM- cell signalling
are often associated with various diseases(85–87).
Cells actively remodel the surrounding ECM rather than passively sensing it. Forces transmitted
through focal adhesions can significantly modify the ECM for physiological functions, including
cancer metastasis and wound healing(88, 89). Matrix remodelling also enables cells to sense distant cells, playing a crucial role in numerous biological processes(90–92).
57
5.6 The Cell Cycle
The mitotic cell cycle consists of two primary phases: interphase and M phase (see Figure 5.3),
which serve to temporally separate the duplication of cellular content in interphase from its subsequent segregation into two genetically identical daughter cells during mitosis. The underlying
regulatory network governing the cell cycle is intricately designed with the specific aim of ensuring the timely and precise duplication and segregation of genomic DNA. Notably, DNA replication,
a fundamental process, takes place during interphase within the S phase, marking the initiation
but not the completion of DNA replication.
Within interphase, the periods distinguishing S phase from M phase have traditionally been
labelled as ‘gap phases’, namely G1 preceding S phase and G2 following S phase. These phases are
not merely temporal intervals within the cell cycle but, rather, they assume critical roles in the regulation of the cell cycle. G1, in particular, holds substantial significance as it serves as the decision
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5 Cellular Structure andFunction
I
M
58
point for cells to either enter the cell cycle or remain in their
Interphase
existing state. During the pre- replicative G1 phase, cells
make the determination to initiate DNA replication and
proceed into the cell cycle. Additionally, cells in G1 possess
G0
S
G1
G2
the option to exit the cell cycle, transitioning into a nonproliferative state known as quiescence or G0.
Crucially, the transition from a non- proliferative state to
G1 represents a prerequisite for cells to initiate DNA replication and embark on the cell cycle. Following the comple-
M
tion of DNA replication, the post- replicative G2 phase
presents another pivotal decision window. In this phase,
cells can commit to entering M phase, characterised by
Mitotic phase
chromatin condensation and the alignment of chromosomes at the cell’s central region. The M phase serves a
Figure5.3 Cell cycle (created in
BioRender).
dual role, ensuring the precise segregation of duplicated
DNA (mitosis) and the division of the entire cellular content into two new daughter cells through cytokinesis.
The M phase effectively commits cells to the segregation of genetic material and initiates the
resetting of the cell cycle for the transition back to interphase(93).
5.7 Control andRegulation ofthe Cell Cycle
The precise regulation of the cell cycle is essential for maintaining DNA integrity in successive
generations and is crucial for proper cell division and proliferation. Aberrant cell cycle regulation
is a hallmark of cancer. Control factors in cell cycle regulation include:
1) Mitogens, which regulate the cell cycle rate by downregulating inhibitory barriers that block
cell cycle progression.
2) Growth factors that control cell growth in terms of size and mass by increasing the synthesis of
proteins and macromolecules needed for the cell cycle.
3) Survival factors that inhibit apoptosis and promote cell survival.
Cell cycle regulation relies on cell cycle checkpoints, which ensure the orderly production of
daughter cells. The three primary checkpoints occur at transitions from G1 to S phase, G2 to M
phase and metaphase to anaphase. These checkpoints monitor growth factor presence, cell size
and DNA damage. A restriction point in late G1marks the transition to growth factor- independent
cell cycle progression. The G2/M checkpoint verifies proper DNA duplication and detects DNA
damage. The metaphase/anaphase checkpoint, also called the spindle apparatus checkpoint,
ensures proper chromosome attachment to the mitotic spindle. Checkpoints are surveillance
mechanisms that involve numerous genes responsible for detecting DNA damage and initiating
cell cycle arrest, DNA repair or apoptosis. The checkpoint surveillance system comprises sensor
proteins, transducers and effectors(94).
5.8 Stem Cells andDifferentiation
Stem cells possess the unique capacity for self- renewal and differentiation into specialised cellular
populations, ultimately forming various tissues and organs in the body. Stem cell differentiation
transitions cells from a non- specific state to a morphologically and functionally specific state(95).
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References
Stem cells fall into two primary categories: embryonic stem cells and somatic stem cells. Totipotent
stem cells can divide and differentiate into cells of the entire organism, forming embryo and
extra- embryonic structures. Pluripotent stem cells give rise to cells of all germ layers but not
extra-embryonic structures. Multipotent stem cells have a narrower differentiation spectrum,
specialising in specific cell lineages. Oligopotent stem cells can differentiate into several cell
types, while unipotent stem cells have limited differentiation capabilities and repeatedly
divide(96). Abnormal stem cell self- renewal and differentiation can lead to human malignancy.
Stem cells participate in various physiological and pathological processes, such as tissue development, wound healing and tumourigenesis(97, 98).
5.9 Summary
The basic structural unit of life is the cell. A thorough understanding of the tissues and organisms that
cells make up is made possible by concentrating on the cell. Understanding the normal structure and
function of the cell is necessary to define a pathological condition. Identification of a pathological
condition will be made easier by defining the normal processes of intracellular connections, cell–cell
communication and signalling pathways, cell- extracellular matrix communication and cell cycle.
Determining the image of the cell’s typical structure and function is crucial for this reason.
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