BOTANY OUTLINE

CHARACTERISTICS OF LIVING THINGS AND MANIFESTATIONS OF LIFE

Life
process

Explanation

Earthworms

Movemen
t

All living things move in some

way. This may be obvious, such
as animals that are able to
walk, or less obvious, such as
plants that have parts that
move to track the movement of
the sun.

Earthworms use circular and

longitudinal muscles to move
through soil or along surfaces.

Respirati
on

Respiration is a chemical
reaction that happens within
cells to release energy from
food.

The food that earthworms eat

supplies their body with energyrich molecules such asglucose. On
entering the cells of their body,
these molecules are broken down
in aseries of steps to release
energy to be used by the body,
producing carbon dioxide and
water as waste products.

Sensitivit
y

The ability to detect changes in

the surrounding environment.

Earthworms have light-sensitive

cells scattered in their outer skin.
Their skin cells are also sensitive
to touch andchemicals.

Growth

All living things grow.

Earthworms hatch from eggs and

can grow up to a metre or more in
length! Some earthworms are
also able to regrow small parts of
their body that have been lost or
injured.

Reproduc
tion

The ability to reproduce and

pass genetic information onto
their offspring.

Earthworms have both sperm and

eggs within their bodies (they are
hermaphrodites) but they cannot
self-fertilise and need to mate
with another individual. After
mating, a cocoon containing the
fertilised eggs is deposited in the
soil.

Excretion

Getting rid of waste.

Earthworms excrete waste from

their anus the last segment of
their body.

Nutrition

The intake and use ofnutrients.

This occurs in very different
ways in different kinds of living
things.

Earthworm nutrition comes from a

variety of sources, depending on
their species. Food types include
manure, compost, plant
material, fungi, microorganisms
and decaying animals. They take
in food through their mouths.

5 MAJOR GROUPS OF LIVING THINGS

Monera (includes Eubacteria and Archeobacteria)
Individuals are single-celled, may or may not move, have a cell wall, have no
chloroplasts or other organelles, and have no nucleus. Monera are usually very tiny,
although one type, namely the blue-green bacteria, look like algae. They are
filamentous and quite long, green, but have no visible structure inside the cells. No
visible feeding mechanism. They absorb nutrients through the cell wall or produce
their own by photosynthesis.
Protista
Protists are single-celled and usually move by cilia, flagella, or by amoeboid
mechanisms. There is usually no cell wall, although some forms may have a cell
wall. They have organelles including a nucleus and may have chloroplasts, so some
will be green and others won't be. They are small, although many are big enough to
be recognized in a dissecting microscope or even with a magnifying glass. Nutrients
are acquired by photosynthesis, ingestion of other organisms, or both.
Fungi
Fungi are multicellular,with a cell wall, organelles including a nucleus, but no
chloroplasts. They have no mechanisms for locomotion. Fungi range in size from
microscopic to very large ( such as mushrooms). Nutrients are acquired by
absorption. For the most part, fungi acquire nutrients from decaying material.
Plantae
Plants are multicellular and most don't move, although gametes of some plants
move using cilia or flagella. Organelles including nucleus, chloroplasts are present,
and cell walls are present. Nutrients are acquired by photosynthesis (they all require
sunlight).
Animalia
Animals are multicellular, and move with the aid of cilia, flagella, or muscular organs
based on contractile proteins. They have organelles including a nucleus, but no
chloroplasts or cell walls. Animals acquire nutrients by ingestion.

A "mini-key" to the five kingdoms

Suppose you see something in freshwater that certainly appears to be living. How
can you begin to determine what it is? Here is a key (not quite perfect) that you
might use to help determine the kingdom to which it belongs.
1. Is it green or does it have green parts?
o

Yes - go to 2

No - go to 3

2. Could be a plant or a protist, or blue-green bacteria. Make sure that the green is
really part of the organism, though. An animal might have eaten something green,
for example.
o

Single-celled? go to 6

Multicellular? Plantae. Look for cell walls, internal structure. In the

compound microscope you might be able to see chloroplasts.

3. Could be a moneran (bacteria), protist, fungus, or animal.

o

Single-celled - go to 4

Multicellular (Look for complex or branching structure, appendages) go to 5

4. Could be a moneran or a protist. Can you see any detail inside the cell?
o

Yes - Protista. You should be able to see at least a nucleus and/or

contractile vacuole, and a definite shape. Movement should be
present, using cilia, flagella, or amoeboid motion. Cilia or flagella may
be difficult to see.

No - Monera. Should be quite small. May be shaped like short dashes

(rods), small dots (cocci), or curved or spiral shaped. The largest them
that is commonly found in freshwater is called Spirillum volutans. It is
spiral shaped, and can be nearly a millimeter long. Except for Spirillum,
it is very difficult to see Monerans except in a compound microscope
with special lighting.

5. Animalia or Fungi. Is it moving?

o

Yes - Animalia. Movement can be by cilia, flagella, or complex,

involving parts that contract. Structure should be complex. Feeding
activity may be obvious.

No - Fungus. Should be branched, colorless filaments. May have some

kind of fruiting body (mushrooms are a fungus, don't forget). Usually
attached to some piece of decaying matter - may form a fuzzy coating
on or around an object. In water, some bacterial infections of fish and
other animals may be mistaken for a fungus.

6. Most likely Protista. If it consists of long, unbranched greenish filaments with no

apparent structure inside, it is blue-green bacteria (sometimes mistakenly called
blue-green algae), a Moneran.
Most green protists are flagellates, that is, they move rapidly with a spiralling
motion. Unless you get them to stop, you can't really see the flagella. Watch out for
colonial protists, though, such as Volvox, which forms a spinning ball of green cells.
Don't be fooled into thinking it is a plant.

CELL REPRODUCTION
Cell Division Functions in Reproduction, Growth, and Repair
Cell division involves the distribution of identical genetic material, DNA, to two
daughters cells. What is most remarkable is the fidelity with which the DNA is
passed along, without dilution or error, from one generation to the next.

Core Concepts:

All Organisms Consist of Cells and Arise from Pre-existing Cells

o

Mitosis is the process by which new cells are generated.

Meiosis is the process by which gametes are generated for

reproduction.

The Cell Cycle Represents All Phases in the Life of a Cell

o

DNA replication (S phase) must precede mitosis, so that all daughter

cells receive the same complement of chromosomes as the parent cell.

The gap phases separate mitosis from S phase. This is the time when
molecular signals mediate the switch in cellular activity.

Mitosis involves the separation of copied chromosomes into separate

cells

Unregulated Cell Division Can Lead to Cancer

o

Cell-cycle checkpoints normally ensure that DNA replication and

mitosis occur only when conditions are favorable and the process is
working correctly.

Mutations in genes that encode cell-cycle proteins can lead to

unregulated growth, resulting in tumor formation and ultimately
invasion of cancerous cells to other organs.

In order to better understand the concept of cell division and genetics, some basic
definitions are in order:

gene - basic unit of heredity; codes for a specific trait

locus - the specific location of a gene on a chromosome (locus - plural loci)

genome - the total hereditary endowment of DNA of a cell or organism

somatic cell - all body cells except reproductive cells

gamete - reproductive cells (i.e. sperm & eggs)

chromosome - elongate cellular structure composed of DNA and protein they are the vehicles which carry DNA in cells

diploid (2n) - cellular condition where each chromosome type is represented

by two homologous chromosomes

haploid (n) - cellular condition where each chromosome type is represented

by only one chromosome

homologous chromosome - chromosome of the same size and shape which

carry the same type of genes

chromatid - one of two duplicated chromosomes connected at the

centromere

centromere - region of chromosome where microtubules attach during

mitosis and meiosis

Chromosome structure

composed
of DNA and
protein
(histones)
all tightly
wrapped
up in one
package

duplicated
chromosom
es are
connected
by a
centromere

Example - an organism is 2n = 4.

Chromosomes 1 & 2 are homologous

chromosomes

Chromosomes 3 & 4 are homologous

chromosomes

Chromosomes 1 & 3 came from the mother

Chromosomes 2 & 4 came from the father

Typical Animal Life Cycle

The Cell Cycle

G1 - first gap
S - DNA
synthesis
(replication)
G2 - second
gap
M - mitosis

mitosis - nuclear/chemical events resulting in two daughter nuclei which

have identical genetic material to each other and to the mother cell

cytokinesis - division of the cytoplasm. This usually occurs with mitosis, but
in some organisms this is not so

Mitosis in a Nutshell

The stages of the cell cycle can be broken down into six stages:
o

Interphase, Prophase, Metaphase, Anaphase, Telophase

Interphase

is the "resting" or non-mitotic portion of the cell cycle.

It is comprised of G1, S, and G2 stages of the cell cycle.

DNA is replicated during the S phase of Interphase

Prophase - the first stage of
mitosis.

The chromosomes
condense and become
visible

The centrioles form and

move toward opposite
ends of the cell ("the
poles")

The nuclear membrane

dissolves

The mitotic spindle forms

(from the centrioles in
animal cells)

Spindle fibers from each

centriole attach to each
sister chromatid at the
kinetochore

Compare Prophase to
the Prophase I and to
the Prophase II stages of
mitosis.

Metaphase

The Centrioles
complete their
migration to the
poles

The chromosomes
line up in the
middle of the cell
("the equator")

Compare Metaphase to
the Metaphase I and to
the Metaphase II stages
of mitosis.
Anaphase

Spindles attached to
kinetochores begin to shorten.

This exerts a force on the

sister chromatids that pulls
them apart.

Spindle fibers continue to

shorten, pulling chromatids to
opposite poles.

This ensures that each

daughter cell gets identical
sets of chromosomes

Compare Anaphase to the Anaphase

I and to the Anaphase II stages of
mitosis.

Telophase

The chromosomes
decondense

The nuclear envelope

forms

Cytokinesis reaches
completion, creating
two daughter cells

Compare Telophase to
the Telophase I and to
the Telophase II stages of
mitosis.

Cytokinesis Divides the Cytoplasm

In animal cells, cytokinesis occurs by a process known as cleavage

First, a cleavage furrow appears

o

A contractile ring of actin microfilaments in association with myosin, a protein

o

cleavage furrow = shallow groove near the location of the old

metaphase plate

Actin and myosin are also involved in muscle contraction and other
movement functions

The contraction of a the dividing cell's ring of microfilaments is like the

pulling of drawstrings
o

The cell is pinched in two

Cytokinesis in plant cells is different because plant cells have cell walls.

There is no cleavage furrow

During telophase, vesicles from the Golgi apparatus move along microtubules
to the middle of the cell (where the cell plate was) and coalesce, producing
the cell plate

Cell-wall construction materials are carried in the vesicles and are

continually deposited until a complete cell wall forms between the two
daughter cells

Chromosome Separation Is the Key Event of Mitosis

Mitotic spindle fibers are the railroad tracks for chromosome movement.
o

Spindle fibers are made of microtubules.

Microtubules are lengthened and shortened by the addition and loss of

tubulin subunits.

Mitotic spindle shortening during anaphase is a result of the loss of

tubulin subunits.

A kinetochore motor is the engine that drives chromosome movement.

o

Multiple studies have shown that the kinetochore contains motor

proteins that can walk along the spindle fiber during anaphase.

These proteins presumably remove tubulin subunits, shortening

spindle fibers and facilitating the chromosome movement.

Regulation of the Cell Cycle

The cell cycle is controlled by a cyclically operating set of reaction sequences that
both trigger and coordinate key events in the cell cycle

The cell-cycle control system is driven by a built-in clock that can be adjusted
by external stimuli (chemical messages)

Checkpoint - a critical control point in the cell cycle where stop and goahead signals can regulate the cell cycle

Animal cells have built-in stop signals that halt the cell cycles and
checkpoints until overridden by go-ahead signals.

Three Major checkpoints are found in the G1, G2, and M phases of the
cell cycle

The G1 checkpoint - the Restriction Point

o

The G1 checkpoint ensures that the cell is large enough to divide, and
that enough nutrients are available to support the resulting daughter
cells.

If a cell receives a go-ahead signal at the G1 checkpoint, it will usually

continue with the cell cycle

If the cell does not receive the go-ahead signal, it will exit the cell cycle
and switch to a non-dividing state called G0

Actually, most cells in the human body are in the G0 phase

The G2 checkpoint ensures that DNA replication in S phase has been

completed successfully.

The metaphase checkpoint ensures that all of the chromosomes are attached
to the mitotic spindle by a kinetochore.

Cyclins and Cyclin-Dependent Kinases - The Cell-Cycle Clock

Rhythmic fluctuations in the abundance and activity of cell-cycle control molecules
pace the events of the cell cycle.

Kinase - a protein which activates or deactivates another protein by

phosphorylating them.

Kinases give the go-ahead signals at the G1 and G2 checkpoints

The kinases that drive these checkpoints must themselves be activated

o

The activating molecule is a cyclin, a protein that derives its name

from its cyclically fluctuating concentration in the cell

Because of this requirement, these kinases are called cyclindependent kinases, or Cdk's

MPF - Maturation Promoting Factor (M-phase promoting factor)

Cyclins accumulate during the G1, S, and G2 phases of the cell cycle

By the G2 checkpoint (the red bar in the figure), enough cyclin is available to
form MPF complexes (aggregations of Cdk and cyclin) which initiate mitosis
o

MPF apparently functions by phosphorylating key proteins in the

mitotic sequence

Later in mitosis, MPF switches itself off by initiating a process which leads to
the destruction of cyclin
o

Cdk, the non-cyclin part of MPF, persists in the cell as an inactive form
until it associates with new cyclin molecules synthesized during
interphase of the next round of the cell cycle

PDGF - Platelet-Derived Growth Factors - An Example of an External Signal

for Cell Division
PDGF is required for the division of fibroblasts which are essential in wound healing

When injury occurs, platelets (blood cells important in blood clotting) release
PDGF

Fibroblasts are a connective tissue cells which possess PDGF receptors on

their plasma membranes

The binding of PDGF activates a signal-transduction pathway that leads to a

proliferation of fibroblasts and a healing of the wound

Density Dependent Inhibition

Cells grown in culture will rapidly divide until a single layer of cells is spread
over the area of the petri dish, after which they will stop dividing

If cells are removed, those bordering the open space will begin dividing again
and continue to do so until the gap is filled - this is known as contact
inhibition

Apparently, when a cell population reaches a certain density, the amount of

required growth factors and nutrients available to each cell becomes
insufficient to allow continued cell growth

Anchorage Dependence

For most animal cells to divide, they must be attached to a substratum, such
as the extracellular matrix of a tissue or the inside of the culture jar

Anchorage is signaled to the cell-cycle control system via pathways involving

membrane proteins and the cytoskeleton

Cells Which No Longer Respond to Cell-Cycle Controls - Cancer Cells

Cancer cells do not respond normally to the body's control mechanism.

o

They divide excessively and invade other tissues

If left unchecked, they can kill the organism

Cancer cells do not exhibit contact inhibition

o

If cultured, they continue to grow on top of each other when the total
area of the petri dish has been covered

They may produce required external growth factor (or override factors)
themselves or possess abnormal signal transduction sequences which
falsely convey growth signals thereby bypassing normal growth checks

Cancer cells exhibit irregular growth sequences

o

If growth of cancer cells does cease, it does so at random points of the

cell cycle

Cancer cells can go on dividing indefinitely if they are given a continual

supply of nutrients

Normal mammalian cells growing in culture only divide 20-50

times before they stop dividing

Meiosis
More definitions:

Allele - alternate forms of the same gene

Homozygous - having two identical alleles for a given gene

Heterozygous - having two different alleles for a given gene

Genotype - genetic makeup of an organism

Phenotype - the expressed traits of an organism

Meiosis in a Nutshell

Meiosis Is a Special Type of Cell Division That Occurs in Sexually Reproducing

Organisms
o

Meiosis reduces the chromosome number by half, enabling sexual

recombination to occur.

Meiosis of diploid cells produces haploid daughter cells, which

may function as gametes.

Gametes undergo fertilization, restoring the diploid number of

chromosomes in the zygote

Meiosis and fertilization introduce genetic variation in three ways:

Crossing over between homologous chromosomes at prophase I.

Independent assortment of homologous pairs at metaphase I:

Each homologous pair can orient in either of two ways at

the plane of cell division.

The total number of possible outcomes = 2n (n = number

of haploid chromosomes).

Random chance fertilization between any one female gamete

with any other male gamete.

The Role of Sexual Reproduction in Evolution

o

Sexual reproduction in a population should decline in frequency

relative to asexual reproduction.

Asexual reproductionNo males are needed, all individuals can

produce offspring.

Sexual reproductionOnly females can produce offspring,

therefore fewer are produced.

Sexual reproduction may exist because it provides genetic variability

that reduces susceptibility of a population to pathogen attack.

The stages of meiosis can be broken down into two main stages, Meiosis
I and Meiosis II

Meiosis I can be broken down into four substages: Prophase I, Metaphase I,

Anaphase I and Telophase I

Meiosis II can be broken down into four substages: Prophase II, Metaphase
II, Anaphase II and Telophase II

Meiosis I

Prophase I - most of the significant processes of

Meiosis occur during Prophase I

The chromosomes condense and become

visible

The centrioles form and move toward the

poles

The nuclear membrane begins to dissolve

The homologs pair up, forming a tetrad

o

Each tetrad is comprised of four

chromotids - the two homologs, each
with their sister chromatid

Homologous chromosomes will swap genetic

material in a process known as crossing
over (abbreviated as XO)
o

Crossing over serves to increase

genetic diversity by creating four
unique chromatids

Compare Prophase I to Prophase II and to

the Prophase stage of mitosis.

Crossing Over

Genetic
material
from
the hom
ologous
chromo
somes is
randomly
swapped

This
creates
four
unique
chromati
ds

Since
each
chromati
d is
unique,
the
overall
genetic
diversity
of the
gametes
is greatly
increase
d

Metaphase I

Microtubules grow from the centrioles and

attach to the centromeres

The tetrads line up along the cell equator

Compare Metaphase I to Metaphase II and to

the Metaphase stage of mitosis.

Anaphase I

The centromeres break and homologous

chromosomes separate (note that
the sister chromatids are still attached)

Cytokinesis begins

Compare Anaphase I to Anaphase II and to

the Anaphase stage of mitosis.

Telophase I

The chromosomes may decondense

(depends on species)

Cytokinesis reaches completion,

creating two haploid daughter cells

Compare Telophase I to Telophase II and to

the Telophase stage of mitosis.

Meiosis II

Prophase II

Centrioles form and move toward the poles

The nuclear membrane dissolves

Compare Prophase II to Prophase I and to the Prophase stage

of mitosis.

Metaphase II

Microtubules grow from the centrioles and attach to the

centromeres

The sister chromatids line up along the cell equator

Compare Metaphase II to Metaphase I and to

the Metaphase stage of mitosis.

Anaphase II

The centromeres break and sister

chromatids separate

Cytokinesis begins

Compare Anaphase II to Anaphase I and to

the Anaphase stage of mitosis.

Telophase II

The chromosomes may decondense

(depends on species)

Cytokinesis reaches completion,

creating four haploid daughter cells

Compare Telophase II to Telophase I and to

the Telophase stage of mitosis.

A Comparison between Mitosis and Meiosis

Some questions to ponder

How does the number of daughter cells produced from mitosis and meiosis
differ?

How does the ploidy of the daughter cells produced from mitosis and meiosis
differ?

Do the daughter cells produced from mitosis contain identical genetic

complements?

Do any of the daughter cells produced from meiosis contain identical genetic
complements?

When do the homologous chromosomes separate during mitosis?

When do the homologous chromosomes separate during meiosis?

When do sister chromatids separate during mitosis?

When do sister chromatids separate during meiosis?

Click the cockroach below to view the answers to these questions.

The Consequences of Meiotic Mistakes

Nondisjunctions occur when homologous chromosomes fail to separate at meiosis I
or when chromatids fail to separate at meiosis II.

Fertilization can result in embryos that are 2n + 1 (a "trisomy") or 2n 1.

Abnormal copy numbers of one or more chromosomes is usually, but not

always, fatal (Example: Down syndrome)

Polyploidy can occur when whole sets of chromosomes fail to separate at meiosis I
or II.

The resulting 2n gametes, if fertilized by normal sperm, create 3n zygotes

(triploid).

Organisms with an odd number of chromosome sets cannot produce viable

gametes (Example: seedless fruits).
PLANT TISSUES

A mature vascular plant (any plant other than mosses and liverworts), contains
several types of differentiated cells. These are grouped together in tissues. Some
tissues contain only one type of cell. Some consist of several.
Meristematic
The main function of meristematic tissue is mitosis. The cells are small, thin-walled,
with no central vacuole and no specialized features.
Meristematic tissue is located in

the apical meristems at the growing points of roots and stems.

the secondary meristems (lateral buds) at the nodes of stems (where

branching occurs) [View], and in some plants,

meristematic tissue, called the cambium, that is found within mature stems
and roots.

The cells produced in the meristems soon become differentiated into one or another
of several types.
Protective
Protective tissue covers the surface of leaves and the living cells of roots and stems.
Its cells are flattened with their top and bottom surfaces parallel. The upper and
lower epidermis of the leaf are examples of protective tissue [View].
Parenchyma
The cells of parenchyma are large, thin-walled, and usually have a large central
vacuole. They are often partially separated from each other and are usually stuffed
with plastids.
In areas not exposed to light, colorless plastids predominate and food storage is the
main function. The cells of the white potato are parenchyma cells.
Where light is present, e.g., in leaves, chloroplasts predominate
and photosynthesis is the main function

Sclerenchyma
The walls of these cells are very thick and built up in a uniform layer around the
entire margin of the cell. Often, the cell dies after its cell wall is fully formed.
Sclerenchyma cells are usually found associated with other cells types and give
them mechanical support.
Sclerenchyma is found in stems and also in leaf veins. [View] Sclerenchyma also
makes up the hard outer covering of seeds and nuts.
Collenchyma

Collenchyma cells have thick walls that are especially thick at their corners. These
cells provide mechanical support for the plant. They are most often found in areas
that are growing rapidly and need to be strengthened. The petiole ("stalk") of
leaves is usually reinforced with collenchyma [View].
Xylem
Xylem conducts water and dissolved minerals from the roots to all the other parts of
the plant.
In angiosperms, most of the water travels in the xylem vessels. These are thickwalled tubes that can extend vertically through several feet of xylem tissue. Their
diameter may be as large as 0.7 mm. Their walls are thickened with secondary
deposits of cellulose and are usually further strengthened by impregnation
with lignin. The secondary walls of the xylem vessels are deposited in spirals and
rings and are usually perforated by pits. [View]
Xylem vessels arise from individual cylindrical cells oriented end to end. At maturity
the end walls of these cells dissolve away, and the cytoplasmic contents die. The
result is the xylem vessel, a continuous nonliving duct.
Xylem also contains tracheids. These are individual cells tapered at each end so
the tapered end of one cell overlaps that of the adjacent cell. Like xylem vessels,
they have thick, lignified walls and, at maturity, no cytoplasm. Their walls are
perforated so that water can flow from one tracheid to the next. The xylem
of ferns and conifers contains only tracheids.
In woody plants, the older xylem ceases to participate in water transport and simply
serves to give strength to the trunk. Wood is xylem. When counting the annual rings
of a tree, one is counting rings of xylem [View].
Phloem
The main components of phloem are

sieve elements and

companion cells.

Sieve elements are so-named because their end walls are perforated. This allows
cytoplasmic connections between vertically-stacked cells. The result is a sieve
tube that conducts the products of photosynthesis sugars and amino acids
from the place where they are manufactured (a "source"), e.g., leaves, to the places
("sinks") where they are consumed or stored; such as

roots

growing tips of stems and leaves

flowers

fruits, tubers, corms, etc.

Sieve elements have no nucleus and only a sparse collection of other organelles.
They depend on the adjacent companion cells for many functions.
Companion cells move sugars, amino acids and a variety of macromolecules into
and out of the sieve elements. In "source" tissue, such as a leaf, the companion
cells use transmembrane proteins to take up by active transport sugars and
other organic molecules from the cells manufacturing them. Water follows
by osmosis. These materials then move into adjacent sieve elements
through plasmodesmata. The pressure created by osmosis drives the flow of
materials through the sieve tubes.
In "sink" tissue, the sugars and other organic molecules leave the sieve elements
through plasmodesmata connecting the sieve elements to their companion cells
and then pass on to the cells of their destination. Again, water follows by osmosis
where it may

leave the plant by transpiration or

increase the volume of the cells or

move into the xylem for recycling through the plant.

Translocation of Food

Food and other organic substances (e.g., some plant hormones and even messenger
RNAs [Link]) manufactured in the cells of the plant are transported in the phloem.

Sugars (usually sucrose),

amino acids, and

other organic molecules

enter the sieve elements through plasmodesmata connecting them to

adjacent companion cells. Once within the sieve elements, these molecules can
be transported either up or down to any region of the plant moving at rates as high
as 110 m per second.
Two demonstrations:

Girdling. Girdling is removing a band of bark from the circumference of the

tree. Girdling removes the phloem but not the xylem. If a tree is girdled in
summer, it continues to live for a time. There is, however, no increase in the
weight of the roots, and the bark just above the girdled region accumulates

carbohydrates. Unless a special graft is made to bridge the gap, the tree
eventually dies as its roots starve.

The photos (courtesy of R. S. Gage and S. Aronoff)

are autoradiographs showing that the products of photosynthesis are
transported in the phloem. A cucumber leaf was supplied with radioactive
water (3HOH) and allowed to carry on photosynthesis for 30 minutes. Then
slices were cut from the petiole of the leaf and covered with a photographic
emulsion. Radioactive products of photosynthesis darkened the emulsion
where it was in contact with the phloem (upper left in both photos), but not
where it was in contact with the xylem vessels (center). In the
photomicrograph on the left, the microscope is focused on the tissue in order
to show the cells clearly; on the right, the microscope has been focused on
the photographic emulsion.

Some fruits, such as the pumpkin, receive over 0.5 gram of food each day through
the phloem. Because the fluid is fairly dilute, this requires a substantial flow. In fact,
the use of radioactive tracers shows that substances can travel through as much as
100 cm of phloem in an hour.
What mechanism drives the translocation of food through the phloem?
Translocation through the phloem is dependent on metabolic activity of the phloem
cells (in contrast to transport in the xylem).

Chilling its petiole slows the rate at which food is translocated out of the leaf
(right).

Oxygen lack also depresses it.

Killing the phloem cells puts an end to it.

The Pressure-Flow Hypothesis

The best-supported theory to explain the movement of food through the phloem is
called the pressure-flow hypothesis.

It proposes that water containing food molecules flows under pressure

through the phloem.

The pressure is created by the difference in water concentration of the

solution in the phloem and the relatively pure water in the nearby xylem
ducts.

At their "source" the leaves sugars are pumped by active transport into
the companion cells and sieve elements of the phloem.

As sugars (and other products of photosynthesis) accumulate in the phloem,

water enters by osmosis.

In the figure, sugar molecules are represented in black, water molecules in

red.)

Turgor pressure builds up in the sieve elements (similar to the creation of root
pressure).

As the fluid is pushed down (and up) the phloem, sugars are removed by the
cortex cells of both stem and root (the "sinks") and consumed or converted
into starch.

Starch is insoluble and exerts no osmotic effect.

Therefore, the osmotic pressure of the contents of the phloem decreases.

Finally, relatively pure water is left in the phloem, and this leaves by osmosis
and/or is drawn back into nearby xylem vessels by the suction
of transpiration-pull.

Thus it is the pressure gradient between "source" (leaves) and "sink" (shoot and
roots) that drives the contents of the phloem up and down through the sieve
elements.
Tests of the theory
1. The contents of the sieve elements must be under pressure.
This is difficult to measure because when a sieve element is punctured with a
measuring probe, the holes in its end walls quickly plug up. However, aphids can
insert their mouth parts without triggering this response.
Left: when it punctures a sieve element, sap enters the insect's mouth parts under
pressure and some soon emerges at the other end (as a drop of honeydew that
serves as food for ants and bees).
Right: honeydew will continue to exude from the mouthparts after the aphid has
been cut away from them. (The photos are the work of the late Martin H.
Zimmerman and were provided by him.)
2. The osmotic pressure of the fluid in the phloem of the leaves must be greater
than that in the phloem of the food-receiving organs such as the roots and fruits.
Most measurements have shown this to be true.
Transport of Messenger RNA (mRNA) through the Phloem
Plant scientists at the Davis campus of the University of California (reported in the
13 July 2001 issue of Science) have demonstrated that messenger RNAs can also
be transported long distances in the phloem. They grafted normal
tomato scions onto mutant tomato stocks and found that

mRNAs synthesized in the stock were transported into the scions.

These mRNAs converted the phenotype of the scion into that of the stock.

Transport of Water and Minerals in Plants

Most plants secure the water and minerals they need from their roots.
The path taken is: soil -> roots -> stems -> leaves
The minerals (e.g., K+, Ca2+) travel dissolved in the water (often accompanied by
various organic molecules supplied by root cells).
Less than 1% of the water reaching the leaves is used in photosynthesis and plant
growth. Most of it is lost in transpiration.
However, transpiration does serve two useful functions:

It provides the force for lifting the water up the stems.

It cools the leaves.

Water and minerals enter the root by separate paths which eventually converge in
the stele.
The Pathway of Water

Soil water enters the root through its epidermis. It appears that water then travels
in both

the cytoplasm of root cells called the symplast that is, it crosses the
plasma membrane and then passes from cell to cell
through plasmodesmata.

in the nonliving parts of the root called the apoplast that is, in the
spaces between the cells and in the cells walls themselves. This water has
not crossed a plasma membrane.

However, the inner boundary of the cortex, the endodermis, is impervious to water
because of a band of lignified matrix called the casparian strip. Therefore, to enter
the stele, apoplastic water must enter the symplasm of the endodermal cells. From
here it can pass by plasmodesmata into the cells of the stele.
Once inside the stele, water is again free to move between cells as well as through
them. In young roots, water enters directly into the xylem vessels and/or tracheids
[link to views of the structure of vessels and tracheids]. These are nonliving
conduits so are part of the apoplast.
Once in the xylem, water with the minerals that have been deposited in it (as well
as occasional organic molecules supplied by the root tissue) move up in the vessels
and tracheids.
At any level, the water can leave the xylem and pass laterally to supply the needs
of other tissues.
At the leaves, the xylem passes into the petiole and then into the veins of the leaf.
Water leaves the finest veins and enters the cells of the spongy and palisade layers.
Here some of the water may be used in metabolism, but most is lost in
transpiration.
The Pathway of Minerals
Minerals enter the root by active transport into the symplast of epidermal cells and
move toward and into the stele through the plasmodesmata connecting the cells.
They enter the water in the xylem from the cells of the pericycle (as well as of
parenchyma cells surrounding the xylem) through specialized transmembrane
channels.
What Forces Water Through the Xylem?
Observations

The mechanism is based on purely physical forces because the xylem vessels
and tracheids are lifeless.

Roots are not needed. This was demonstrated over a century ago by a
German botanist who sawed down a 70-ft (21 meters) oak tree and placed
the base of the trunk in a barrel of picric acid solution. The solution was
drawn up the trunk, killing nearby tissues as it went.

However, leaves are needed. When the acid reached the leaves and killed
them, the upward movement of water ceased.

Removing a band of bark from around the trunk a process

called girdling does not interrupt the upward flow of water. Girdling
removes only the phloem, not the xylem, and so the foliage does not wilt. (In
due course, however, the roots and thus the entire plant will die
because the roots cannot receive any of the food manufactured by the
leaves.)

Transpiration-Pull
In 1895, the Irish plant physiologists H. H. Dixon and J. Joly proposed that water is
pulled up the plant by tension (negative pressure) from above.
As we have seen, water is continually being lost from leaves by transpiration. Dixon
and Joly believed that the loss of water in the leaves exerts a pull on the water in
the xylem ducts and draws more water into the leaf.
But even the best vacuum pump can pull water up to a height of only 34 ft (10.4 m)
or so. This is because a column of water that high exerts a pressure of ~15
lb/in2 (103 kilopascals, kPa) just counterbalanced by the pressure of the
atmosphere. How can water be drawn to the top of a sequoia (the tallest is 370 feet
[113meters] high)? Taking all factors into account, a pull of at least 270 lb/in 2 (~1.9
x 103 kPa) is probably needed.
The answer to the dilemma lies the cohesion of water molecules; that is the
property of water molecules to cling to each through the hydrogen bonds they form.
When ultrapure water is confined to tubes of very small bore, the force of cohesion
between water molecules imparts great strength to the column of water. It has been
reported that tensions as great as 3000 lb/in 2 (21 x 103 kPa) are needed to break the
column, about the value needed to break steel wires of the same diameter. In a
sense, the cohesion of water molecules gives them the physical properties of solid
wires.
Because of the critical role of cohesion, the transpiration-pull theory is also called
the cohesion theory.
Some support for the theory

If sap in the xylem is under tension, we would expect the column to snap
apart if air is introduced into the xylem vessel by puncturing it. This is the
case.

If the water in all the xylem ducts is under

tension, there should be a resulting inward
pull (because of adhesion) on the walls of

the ducts. This inward pull in the band of sapwood in an actively transpiring
tree should, in turn, cause a decrease in the diameter of the trunk.
The graph shows the results of obtained by D. T. MacDougall when he made
continuous measurements of the diameter of a Monterey pine. The diameter
fluctuated on a daily basis reaching its minimum when the rate of
transpiration reached its maximum (around noon)

The rattan vine may climb as high as 150 ft (45.7 m) on the trees of the
tropical rain forest in northeastern Australia to get its foliage into the sun.
When the base of a vine is severed while immersed in a basin of water, water
continues to be taken up. A vine less than 1 inch (2.5 cm) in diameter will
"drink" water indefinitely at a rate of up to 12 ml/minute.
If forced to take water from a sealed container, the vine does so without any
decrease in rate, even though the resulting vacuum becomes so great that
the remaining water begins to boil spontaneously. (The boiling temperature of
water decreases as the air pressure over the water decreases, which is why it
takes longer to boil an egg in Denver than in New Orleans.)

Transpiration-pull enables some trees and shrubs to live in seawater.

Seawater is markedly hypertonic to the cytoplasm in the roots of the red
mangrove (Rhizophora mangle), and we might expect water to leave the cells
resulting in a loss in turgor and wilting. In fact, the remarkably high tensions
(on the order of 500800 lb/in2[~3 to 5 thousand kPa]) in the xylem can pull
water into the plant against this osmotic gradient. So mangroves literally
desalt seawater to meet their needs.

Problems with the theory

When water is placed under a high vacuum, any dissolved gases come out of
solution as bubbles (as we saw above with the rattan vine). This is
called cavitation. Any impurities in the water enhance the process. So
measurements showing the high tensile strength of water in capillaries require
water of high purity not the case for sap in the xylem.
So might cavitation break the column of water in the xylem and thus interrupt its
flow? Probably not so long as the tension does not greatly exceed 270 lb/in 2 (~1.9 x
103 kPa).
By spinning branches in a centrifuge, it has been shown that water in the xylem
avoids cavitation at negative pressures exceeding 225 lb/in 2 (~1.6 x 103 kPa). And
the fact that sequoias can successfully lift water 358 ft (109 m) which would
require a tension of 270 lb/in2 (~1.9 x 103 kPa) indicates that cavitation is avoided
even at that value.

However, such heights may be approaching the limit for xylem transport.
Measurements close to the top of the tallest living sequoia (370 ft [=113 m] high)
show that the high tensions needed to get water up there have resulted in:

smaller stomatal openings, causing

lower concentrations of CO2 in the needles, causing

reduced photosynthesis, causing

reduced growth (smaller cells and much smaller needles).

(Reported by Koch, G. W. et al., in Nature, 22 April 2004.)

So the limits on water transport limit the ultimate height which trees can reach. The
tallest tree ever measured, a Douglas fir, was 413 ft. (125.9 meters) high.
Root Pressure
When a tomato plant is carefully severed close to the base of the stem, sap oozes
from the stump. The fluid comes out under pressure which is called root pressure.
Root pressure is created by the osmotic pressure of xylem sap which is, in turn,
created by dissolved

minerals and

sugars

that have been actively transported into the apoplast of the stele.
One important example is the sugar maple when, in very early spring, it hydrolyzes
the starches stored in its roots into sugar. This causes water to pass by osmosis
through the endodermis and into the xylem ducts. The continuous inflow forces the
sap up the ducts.
Although root pressure plays a role in the transport of water in the xylem in some
plants and in some seasons, it does not account for most water transport.

Few plants develop root pressures greater than 30 lb/in 2 (207 kPa), and some
develop no root pressure at all.

The volume of fluid transported by root pressure is not enough to account for
the measured movement of water in the xylem of most trees and vines.

Those plants with a reasonably good flow of sap are apt to have the lowest
root pressures and vice versa.

The highest root pressures occur in the spring when the sap is
strongly hypertonic to soil water, but the rate of transpiration is low. In

summer, when transpiration is high and water is moving rapidly through the
xylem, often no root pressure can be detected.
So although root pressure may play a significant role in water transport in certain
species (e.g., the coconut palm) or at certain times, most plants meet their needs
by transpiration-pull.