Cytoskeleton

• present in all eukaryotic cells.

• 3-dimensional, interconnected network of fibrous protein.
• Components:
1. Microtubules – long rods (~24 nm in diam.), assembled
from subunits of a globular protein called tubulin, which is
made up of 2 polypeptides (α-tubulin and β-tubulin).
The microtubules are formed when tubulin subunits
spontaneously self-assemble into long chains called
protofilaments. Protofilaments line up laterally to form the
microtubule wall.
2. Microfilaments – solid threads composed of actin, also a
globular protein. A microfilament is composed of 2 parallel
chains of actin twisted to form a helix.
3. Intermediate Microfilaments (little is known in plant cells;
more common in animal cells).
(A) Microtubule (B) Microfilament
Cell Plate (Phragmoplast) Formation
in Cytokinesis
• Cytoplasmic streaming – continuous flow of
cytoplasmic particles and organelles around the
periphery of the cell  involves the formation of
microfilament bundles parallel to the direction of
cytoplasmic flow.
• Tip growth – example: pollen tube develops a tubular
extension that grows down the stigma of the flower and
serves to deliver the male nucleus to the egg.
Microtubules guide the cell wall precursors through the
cytoplasm to the growing tip.
Organelles Unique to Plants –
Plastids & Vacuoles
 Plastids
• double membrane-bound organelles in plants.
• contain their own DNA (in nucleoid region) and ribosomes.

• semi-autonomous and reproduce by fission similar to the division

process in prokaryotes.

If plastids only arise from other plastids and can’t be built "from
scratch", then where do they come from?
The egg. Plastids are inherited cytoplasmically, primarily through the
female. (However, there are examples of paternal inheritance of
plastids.) The plastid DNA carries several genes (e.g., large subunit
of rubisco, and genes for resistance to some herbicides.

• Chemistry of the membranes differs from the plasma membrane -

plastid membranes are comprised of glycosylglycerides rather than
phospholipids (phosphate in the polar head group in
glycosylglycerides is replaced with galactose or a related sugar).
 Types of plastids

1. Proplastids - small, precursors to the other plastid

types, found in young cells, actively growing tissues;

2. Chloroplasts - sites of photosynthesis (energy capture).

They contain photosynthetic pigments including
chlorophyll, carotenes and xanthophylls. The chloroplast
is packed with membranes, called thylakoids. The
thylakoids may be stacked into pancake- like piles called
grana (granum, singular). The "liquid" material in the
chloroplast is the stroma. A chloroplast is from 5-20 μm
in diameter and there are usually 50-200 per cell. The
chloroplast genome has about 145 kbase pairs, it is
smaller than that of the mitochondria (200
kbases). About 1/3 of the total cell DNA is extranuclear
(in the chloroplasts and mitochondria).
3. Chromoplasts - non-photosynthetic, colored
plastids; give some fruits (tomatoes, carrots) and
flowers their color;

4. Amyloplasts - colorless, starch-storing plastids.

(leucoplast - another term for amyloplast)

6. Etioplasts - plastids whose development into

chloroplasts has been arrested (stopped). These
contain a dark crystalline body, prolamellar
body, which is essentially a cluster of thylakoids
in a somewhat tubular form.
Plastids can dedifferentiate and convert
from one form into another. For example,
think about the ripening processing in
tomato. Initially, green tomatoes have
chloroplasts which then begin to
accumulate lycopene (red) and become
chromoplasts.

Usually you find only chromoplasts or

chloroplasts in a cell, but not both.
 Vacuoles

• This is the large, central cavity containing fluid,

called cell sap, found in plant cells. The
vacuole is surrounded by a membrane
(tonoplast).
• The vacuole is penetrated by strands of
cytoplasm - transvacuolar strands.
• The tonoplast and plasma membrane have
different properties, such as thickness (tonoplast
thicker).
• The vacuole makes up to 90% or more of the cell
volume. Meristematic and embryonic cells are
exceptions. Young tissues have many small vacuoles
(provacuoles). As the cell grows the vacuoles expand
and eventually coalesce. These small vacuoles appear
to be derived from the Golgi.

• central vacuole -- contains water, ions, organic acids,

sugars, enzymes, and a variety of secondary
metabolites.

• lytic vacuoles -- contain digestive or hydrolytic

enzymes, e.g., proteases (digest protein), ribonucleases
(digest RNA) and glycosidases (break links between
monosaccharides). These enzymes are typically not
used for recycling cellular components
but rather leak out on cell senescence.

• protein bodies -- store proteins.

Lytic Vacuoles and Protein Bodies
 Why do plants cells have a
large central vacuole?

Important roles of the vacuoles:

• A. Energetically efficient means to increase surface

to volume ratio in the dendritic growth form

Since 90% of the cell volume is vacuole, therefore

~90% of the cell is water, which is relatively cheap in
metabolic terms. Thus, it allows plants to get big with a
minimal energy investment.

Plants are particularly 'smart,' since, the cell wall,

which comprises much of the remaining 10% or so of the
cell is a polymer of glucose.
 Cellulose is a great bargain! It is stronger and
cheaper than comparable polymers.

compound/production tensile strength

value (newton m2 x 109)
polymer (weight product/weight
glucose to make)

cellulose 0.83 30

collagen 0.40 2

silk 0.40 10
• B. Water storage
Probably a minor role; mostly in succulents

• C. Waste disposal
- the vacuole is analogous to the lysosome.

• D. pH regulation
The vacuole is a pool to dump excess
protons. There is an active proton pump in the
tonoplast. The cell sap has a pH of 2-5.7,
whereas the cytosol is ca. 7.0.
• E. Storage of essential ions
Ions are pumped into the vacuole for water
balance. Potassium and calcium in particular,
are stored in the vacuole.

• F. Cell enlargement
Cell growth requires some force to allow for
the cell to increase in size. Water pressure
provides the force and it moves into the vacuole.
For example, root hair enlargement is due
entirely to vacuolar enlargement.
• G. Facilitates diffusion
The cytosol essentially forms a thin coating around the
large vacuole which in effect, increases the surface-to-
volume ratio of the cytoplasm. It provides easier access
and shorter diffusion distances between any part of the
cytoplasm and the external environment of the cell. This
can be particularly important for chloroplasts.

• H. Structural support
The vacuole helps to maintain turgor pressure in plant
cells due to the opposing forces of tensile strength of the
wall vs. compression strength of water.
 Plasmodesmata Interconnect Living Plant Cells

• membrane-lined channels traversing the cell wall and connect the cytoplasms
of adjacent cells forming a continuum (symplast).
 cell-cell communication
• contain cytoplasm, plasma membrane, and desmotubule.