3.

Plant cell biology and growth regulators

1. Plant cell, cell wall, and tissue types

2. Cell division, growth, differentiation and

senescence

3. Cell fate and pattern formation in plants

4. Phytohormones: types and functions

Structurally, plant and animal cells are very similar because they are both eukaryotic
cells. They both contain membrane-bound organelles such as the nucleus, mitochondria,
endoplasmic reticulum, golgi apparatus, lysosomes, and peroxisomes.

Both also contain similar membranes, cytosol, and cytoskeletal elements. The functions of
these organelles are extremely similar between the two (peroxisomes perform additional
complex functions in plant cells).

However, the few differences that exist between plant and animals are very significant and
reflect a difference in the functions of each cell.
 Differences
Larger than animal cells (10-30μm Vs 10-100μm)

Cell wall: rigid cell wall (0.1-10 μm) surrounding the cell membrane. Gives added
stability and protection

Chloroplasts: large, double membrane-bound structures (~5 μm across) that contain

chlorophyll, which absorbs sunlight for photosynthesis (up to 100 nos/cell)

Vacuoles: large, liquid-filled organelles found only in plant cells. Vacuoles can occupy
up to 90% of a cell's volume and have a single membrane. Contain a number of
enzymes that perform diverse functions, and their interiors can be used as storage for
nutrients or, provide a place to degrade unwanted substances. (in animal cells if found
they are small)

Peroxisomes: Breakdown of very long chain fatty acids into CO2 and water (beta-
oxidation). In animal cells, the long fatty acids are converted to medium chain fatty
acids, which are subsequently shuttled to mitochondria where they are eventually
broken down to carbon dioxide and water. In yeast and plant cells, this process is
carried out exclusively in peroxisomes.

Centriole: absent (a pair of minute cylindrical organelles near the nucleus in animal
cells, involved in the development of spindle fibers in cell division)
 Cell walls
A rigid, semi-permeable layer of polysaccharides lying outside the plasma membrane of
the cells of plants
Composed of polysaccharides (cellulose, hemicellulose and pectin), cell wall proteins and
phenolic compounds (lignin).

Cell wall largely determines the size and shape of the cell. Provides the strength and
flexibility needed to allow for rapid cell growth

Contains variety of enzymes and play crucial roles in absorption, transport and
secretion of substances.

Also involved in various signal transduction process and defense against pathogens.

Cell wall composition varies

depending on the organism.
 Cell wall structure
The plant cell wall is multi-layered and consists of up to three sections.

Middle lamella: outer cell wall layer that

contains polysaccharides called pectins
(cements CWs of adjacent cells together).

Primary cell wall: layer formed between the

middle lamella and plasma membrane in
growing plant cells.

Primarily composed of cellulose microfibrils

contained within a gel-like matrix of
hemicellulose fibers and pectin
polysaccharides.

Secondary cell wall: This layer is formed

between the primary cell wall and plasma
membrane in some plant cells. Once the Cellulose microfibrils embedded in a matrix
primary cell wall has stopped dividing and of hemicelluloses, pectin and glycoproteins.
growing, it may thicken to form a secondary Often also contains liginin, cutin, waxes etc
cell wall. This rigid layer strengthens and
supports the cell. In addition to cellulose and
hemicellulose, some secondary cell walls
contain lignin.
 Cell wall components
Lignin: a complex polymer of monolignols (aromatic alcohols) (support)

Pectin: a heteropolysaccharide (polymeric carbohydrates) (jelling- hydrophilic)

Cellulose: a polysaccharide (C6H10O5)x of glucose units (Principal component)

Hemicelluose: any plant cell wall polysaccharides, other than cellulose and pectin
(arabinoxylans, xyloglucan etc) (tethering agent)

Callose: β-1,3-glucan polysaccharide (deposited in response to wounding)

Cutin: waxy substance found in the epidermal cell wall (reduce water loss)

Suberin: waxy substance found in the cork cell wall (reduce water loss)
 Cell wall functions
Support: The cell wall provides mechanical strength and support. It also controls the
direction of cell growth.

Withstand turgor pressure: Turgor pressure is the force exerted against the cell wall
as the contents of the cell push the plasma membrane against the cell wall. This
pressure helps a plant to remain rigid and erect, but can also cause a cell to rupture.

Regulate growth: The cell wall sends signals for the cell to enter the cell cycle in
order to divide and grow.

Regulate diffusion: The cell wall is porous allowing some substances, including
proteins, to pass into the cell while keeping other substances out.

Communication: Cells communicate with one another via plasmodesmata (pores or

channels between plant cell walls that allow molecules and communication signals to
pass between individual plant cells).

Protection: The cell wall provides a barrier to protect against plant viruses and other
pathogens. It also helps to prevent water loss.

Storage: The cell wall stores carbohydrates for use in plant growth, especially in
seeds.
The cell wall has an important function in regulating how plant cells achieve their final siz
consequently have an essential role in regulating plant growth.
 Cellulose
In flowering plants, the glucose polymer cellulose is assembled into long microfibrils
a few nanometers in diameter. The rigidity and orientation of these microfibrils control
cell expansion; therefore, cellulose synthesis is a key factor in the growth and
morphogenesis of plants

Each cellulose microfibril is an array of about two to three dozen (1→4)-β-D-glucan

chains (that is 24 to 36 glucan chains are assembled into a functional microfibril).

Cellulose is made up of repeating

monomers of glucose attached end
β-glucans are chains of D-glucose
to end.
polysaccharides linked by β-type glycosidic bonds
Cellulose polymers are bundled into
microfibrils (10-25nm).
 particle rosettes

The microfibrils are synthesized at the plasma membrane by complexes of six-

membered “particle rosettes” that produce a single microfibril
Particle rosettes (cellulose synthase complexes) are six-membered hexagonal arrays of
an estimated six cellulose synthases (CesAs) each.

Rosettes are often located at the ends of cellulose microfibrils

Cellulose synthase (CesA) proteins are integral membrane proteins, approximately

1,000 amino acids in length
Cellulose microfibril synthesis

Cellulose synthesis proceeds by the attachment of glucosyl residues to the growing

terminus of the acceptor glucan chain.

Each CesA unit in the CSC complex synthesizes a single glucan chain
 Cell expansion and Organ growth

Cell expansion requires the

synthesis of new cell wall material
and controlled loosening of the
wall to allow it to stretch and
increase in area.

Cell-wall-associated proteins of the

so-called expansin family are key
components in this process.

Expansins increase wall extensibility,

presumably by breaking non-covalent
bonds between wall polysaccharides
and allowing them to slide relative to
each other

Auxin also plays a key role in cell expansion

 Acid growth hypothesis
Acid growth refers to the ability
of plant cells and plant cell walls
to elongate or expand quickly at
low (acidic) pH

The plant cell wall has high tensile

strength and must be loosened to
enable the cell to grow.

The growth of a plant cell is primarily driven by the uptake of water into the cytoplasm
and vacuole of the plant cell
Cell growth
 Plasmodesmata
Microscopic channels (50–60 nm in diameter), which traverse the cell walls of plant
cells and some algal cells, enabling transport and communication between them.

May have between 103 and 105 plasmodesmata per cell and are constructed of three
main layers, the PM, the cytoplasmic sleeve, and the desmotubule.

The desmotubule is a tube of flattened endoplasmic

reticulum that runs between two adjacent cells.

There are two forms of plasmodesmata: primary

plasmodesmata, which are formed during cell division,
and secondary plasmodesmata, which can form between
mature cells

 Size exclusion limit: highly variable and is

subject to active modification (MP-30 is able to
increase the size exclusion limit from 700
Daltons to 9400)

Transport proteins (TFs), short interfering RNA, messenger RNA, viroids, and viral
genomes from cell to cell.

Transport is mediated by interactions with proteins localized on the desmotubule,

and/or by chaperones partially unfolding proteins, allowing them to fit through the
narrow passage
Chloroplast
Type of plastid— a round, oval, or disk-shaped body

Chloroplasts are distinguished from other types of plastids by their green color, which
results from the presence of two pigments, chlorophyll a and chlorophyll b.

In plants, chloroplasts occur in all green tissues, though they are concentrated
particularly in the leaf mesophyll cells

The chloroplasts are only semiautonomous. They

lack a cell wall and have a reduced prokaryotic type
of genome, which is present in multiple copies. The
chloroplast stroma with its thylakoids is the homolog
of the cyanobacterial cytoplasm but, although
photosynthesis is maintained as the principal
metabolic function, many other functions have been
lost.

The chloroplasts are enclosed within an envelope of

two membranes. The chloroplast stroma contains
strands of prokaryotic-type DNA and ribosomes
that are of similar size (70S) to those in cyanobacteria
but smaller than those found in the cytoplasm of
eukaryotes (80S).
 The chloroplasts are semiautonomous
The size of chloroplast genomes ranges from 120 to 160 kb (in land plants)
The size of prokaryotic ancestor (cyanobacteria) genomes is at least 20–30 times
larger than plant chloroplast genomes

Chloroplast genome
The chloroplast genome most
commonly includes around 100
genes

Chloroplast division
is regulated by the
cell cycle so as to
occur only once, in
the S phase

Over time, many parts of the chloroplast genome were transferred to the nuclear
genome of the host a process called endosymbiotic gene transfer.
 Chloroplast
Chloroplasts are roughly 1–2 μm (1 μm =
0.001 mm) thick and 5–7 μm in diameter.

Enclosed in a chloroplast envelope, which consists of a double membrane with outer

and inner layers, between which is a gap called the intermembrane space.

A third, internal membrane, extensively folded and characterized by the presence of

closed disks (or thylakoids), is known as the thylakoid membrane.

In most higher plants, the thylakoids are arranged in tight stacks called grana
(singular granum). Grana are connected by stromal lamellae, extensions that run
from one granum, through the stroma, into a neighbouring granum.

The thylakoid membrane envelops a central aqueous region known as the thylakoid
lumen.

The space between the inner membrane and the thylakoid membrane is filled with
stroma, a matrix containing dissolved enzymes, starch granules, and copies of the
chloroplast genome.
Chloroplast division
Vacuoles

Vacuole is a sac containing a liquid cell sap surrounded by a membrane (tonoplast)-

mainly involved in regulating the movements of ions around the cell

Temporary storage of materials. Some also store waste products (so the rest of the cell
is protected from contamination)

Help to maintain cell shape and homeostasis.

 Turgor pressure

Aside from storage, the main role of the central vacuole is to maintain turgor pressure
against the cell wall.
Proteins found in the tonoplast (aquaporins) control the flow of water into and out of
the vacuole through active transport, pumping potassium (K+) ions into and out of the
vacuolar interior.
Due to osmosis, water will diffuse into the vacuole, placing pressure on the cell wall. If
water loss leads to a significant decline in turgor pressure, the cell will plasmolyze.

Turgor pressure exerted by vacuoles is also required for cellular elongation: as the cell
wall is partially degraded by the action of expansins, the less rigid wall is expanded by
the pressure coming from within the vacuole. Turgor pressure exerted by the vacuole
is also essential in supporting plants in an upright position.

Another function of a central vacuole is that it pushes all contents of the cell's
cytoplasm against the cellular membrane, and thus keeps the chloroplasts closer to
light.
 Cell wall biosynthesis: key events
 Cellulose is the major
polysaccharide found in
plants. consists of long parallel
linear β-1,4-D-glucan chains
assembled into microfibrils by
hydrogen bonds.

 Synthesis occurs in the

plasma membrane, catalyzed
by cellulose synthases (CESAs)
(Glycosyltransferase Family 2)

CSCs are large multisubunit

complexes containing at least
three different cellulose
synthase enzymes and other
proteins- visualized as rosettes
 Rosette structures transfer Glc
from cytosolic UDP-Glc to
produce multiple extracellular glucan chains that eventually coalesce into a cellulose microfibril.

The current model for cellulose synthesis catalyzed by CESAs: CESA complex takes
the substrate (glucose donor uridine diphosphate (UDP)-Glc) from the cytosol and
transformed into an elongating cellulose polymer. This is then exported across the
plasma membrane.
 Cell wall biosynthesis

Hemi‐celluloses (HC) and pectin are synthesised and modified in the endomembrane
system before being deposited into the apoplast for assembly into the cell wall.

HC is synthesized by a set of CESA-related cellulose synthase-like genes (CSLs). Being

H-boned to cellulose microfibrils- HC limit the extensibility of the cell wall.

Lignin monomers are synthesised in the cytoplasm and transported to the wall.

Laccases have an essential role in lignin formation by catalysing the oxidative coupling
of the lignin monomers in the apoplast.

These individual components are then cross‐linked and assembled into the complex
matrix structure that comprises functional cell walls.

Cell wall composition varies greatly between cell types and can change as the cell
differentiates and with the developmental stages of cells and the plant.

~2000 genes are required for the synthesis and metabolism of cell wall components and
involves multiple cellular compartments.