Gas exchange
CO2 in for the process of photosynthesis
O2 in or out, depending on diffusion gradients. Oxygen is used in respiration in every
living cell, and is a by-product of photosynthesis
Tissues of the Leaf (Anatomy)
(surface to surface)
Leaf Epidermis
- Covers leaf surface
- Generally one cell thick
- Functions
- Covered with cuticle
(thickness varies)
Special cells in leaf epidermis
- Guard cells
which form stomata
- Concentrated on lower epidermis in most leaves
- Pairs of "bean-shaped" cells with thick, rigid inner walls in dicots and most monocots
- Contain chloroplasts
- Opening between guard cells forms stomata
for gas exchange
- Stomata open and close with changes in turgor
- Thin walls stretch as cell swells with water and creates stomate (thick walls don't)
- Photosynthesis and osmotic changes are necessary for most stomate function.
- Epidermal hairs (trichomes)
Silky, woolly, prickly, felt-like, scaly, etc.
There are keys for identifying different types of hairs
Unicellular or multicellular
Glandular (containing stinging chemicals, aromatic oils or crystals)
Leaf Mesophyll
- All internal cells of the leaf outside of vascular bundles
- Parenchyma cells
- Location of Photosynthesis
- Types of Mesophyll
Dicot Leaf Mesophyll
- Palisade mesophyll
- Located near upper epidermis
- Cells elongated to surface
- Contain many chloroplasts
- Spongy mesophyll
- Located near lower epidermis
- Many air spaces
- Isodiametric
- Important in transpiration and CO2 movement
Monocot Leaf Mesophyll
- Monocots usually do not have a distinctive palisade and spongy mesophyll.
- Monocots have parallel veins with a general mesophyll of loosely packed parenchyma
cells on both sides of the veins extending to the epidermis layers.
Vascular Tissue in Leaves (Veins)
Provides support Veins branch extensively throughout the mesophyll of leaves.
- Conduction
- Xylem
(Toward upper epidermis)
Vessels
Fibers
- Phloem
(Toward lower epidermis)
Sieve tubes
Companion cells
Transfer cells
- Bundle sheath
of fibers or parenchyma
- Border parenchyma at tips of "veinlets"
- The leaf petiole
is primarily a vascular connection from the stem to leaf blade.
Vein Patterns
In dicots
, the veins form a network throughout the mesophyll, branching from the mid vein.
In a leaf cross section, the large mid vein is conspicuous, with bundle sheath extensions
which often go from epidermis to epidermis layer, providing additional support.
There may be some secondary growth in the mid vein. Branching veins may be seen
either in cross section or in longitudinal section. Larger branching veins may also
have bundle sheath extensions.
In monocots
, the parallel veins are seen in cross section, along with the cross section of the
leaf, and are more or less uniform in size and distribution across the leaf's interior.
Some grasses may have large thin-walled cells along both sides of the midvein in the
upper epidermis. These cells, called bulliform cells
, help the leaf to fold or roll inward during water deficit periods. This folding
of the leaf probably minimizes evaporation.
In addition, many monocots have enlarged bundle sheath cells, surrounding the veins,
which have a function in photosynthesis for some plants.
Some monocots also have variations in guard cell structure
. Rather than having thickened inner walls, the walls of the guard cells of some
monocots are thickened at the ends, so the guard cells have more of a "dumbbell shape
than a bean shape. The guard cells are also associated with specialized adjacent
cells, called subsidiary cells.
Some Leaf Variations
Modifications for Habitat
Hydromorphic Leaves
- Floating Leaves
- Floating leaves will have stomata on the upper epidermis with air channels into the
palisade parenchyma.
- The spongy mesophyll will be filled with huge air spaces (aerenchyma)
- Vascular tissue may be reduced, especially in the amount of xylem
in the branching veins.
Submerged Leaves
Plants which grow completely underwater will typically have very dissected leaves,
or narrow linear leaves. These shapes minimize resistance to water currents, and
probably prevent damage to tissues which would be more likely with a broad surface.
Some aquatic plants produce dissected leaves where submerged, and less dissected shapes
when the leaves are produced above the surface of the water.
Xeromorphic Leaves
Plants which live in arid environments are subject to drought, and often, intense
sunlight. Such plants are called xerophytes. These plants are subjected to intense
evaporation of water, a resource which is often in short supply. Many such plants
have a number of leaf modifications to prevent excessive water loss.
- Thick cuticle
- The upper epidermis
may be several layers thick.
- The palisade parenchyma
, beneath the epidermis layers may also be thickened.
- Veins
may have bundle sheath extensions in additional to the bundle sheath layer.
- The lower epidermis
may have several layers and a thickened cuticle.
- Deep invaginations of the lower epidermis layer, called stomatal crypts
, are common. Crypts often have many hairs, which in the depression of the crypt
create a micro-environment which is more humid. Stomata are located in the crypts.
Some xeromorphic plants may have succulent leaves, and may have stomata which open
at night rather than in the daylight, again to minimize water loss.
Conifer Leaves (Needles)
The needles of conifers have many of the same adaptations of xeromorphic leaves.
Conifers predominate in areas that have long, cold winters and dryer summers. To
survive conifers are adapted to conditions of moisture stress. A typical conifer
needle will have sunken stomata
, often in channels on the lower surface, thick cuticle
, hypodermis
layers, and, in addition, an endodermis
surrounding the vein (or rarely, veins). As with other conifer organs. the leaves
will have resin canals, lined with parenchyma.
Other Leaf Variations
Shade/Sun
Leaves in the shade exhibit etiolation (small leaves, long stems, thinner blades,
etc.)
Age
In some plants, the shape of leaves changes from the juvenile regions of the plant
to those which are more mature. English ivy leaves show these changes
Leaves and Water Relations in Plants
Stomata are open in daytime which permits diffusion of CO2 into the leaf for photosynthesis. At the same time water is lost through the stomata
by a process called transpiration.
The intercellular spaces of plant tissues are near 100% humidity, and the stomata
are openings into the environment, which is usually not at 100% humidity. The diffusion
gradient for water is from the leaf to the environment. This creates serious problems for water maintenance. The resolution of this problem is
the closure of stomata at night so that water loss is restricted to the daytime hours
when the plant is actively using CO2. It is important to remember that the primary function of stomata is gas exchange,
a subject that will be discussed later.
Transpiration also plays a role in the movement of water throughout the plant as we
shall also discuss later. Transpiration loss is significant. In corn fields, as
much as 90% of the water absorbed by the roots is lost by transpiration.
Other Water Movement - Positive Root Pressure
Simple diffusion pressure in roots moves H2O upward - often forcing the H2O to be exuded from vein tips in leaves, a phenomenon called guttation. The special leaf tip cells are called hydathodes.
Guttation occurs when there is high soil moisture and low evaporation stress. You can typically see guttation in Seattle during the spring and early summer, in the early mornings before the sunlight evaporates the water. Grasses and many herbaceous plants, such as strawberries. guttate nicely. Guttation can not be observed during rain, since the rain drops coat the plant surfaces, and should not be confused with dew, which can condense from the atmosphere onto plant surfaces.
Leaf Abscission
(Loss of leaves)
Loss and replacement of leaves in plants is a normal process. A perennial which loses
all of its leaves at one time in response to seasonal or climate differences, is
termed deciduous
. All leaves, however, have a finite life span, and are lost from the plant, to be
replaced by newer leaves produced in the shoot meristems. The process of leaf loss
is called abscission
, and is controlled by hormones.
The abscission zone is located at the base of the petiole in a region of undifferentiated,
small parenchyma cells. Their walls contain no lignin, and the vascular cells in
the abscission zone are also reduced in size.
The process of abscission is initiated and proceeds as follows:
- The parenchyma cells start dividing rapidly.
- They secrete a layer of suberin in the walls nearest the stem
- The middle lamella, cell walls and cells of the abscission zone dissolve (enzymatic
degradation)
- Leaf abscises
The hormones (which will be discussed later) involved in leaf abscission are:
Auxin
ABA (Abscissic acid)
Ethylene
Associated with abscission, and occurring prior to the loss of the leaf, is a process
which involves degrading and moving a number of solutes, minerals and other substances
from the leaf tissue into the stem and roots. The degradation of chlorophyll and
change in leaf color which results, is often referred to as "fall color". As chlorophyll
degrades the natural carotenoids become prominent. In addition, some leaves may
accumulate anthocyanins in their vacuoles, adding reds or bluish reds to the fall
color. Photoperiod is important in triggering the fall change in pigmentation.
Some Leaf Modifications
(Discussed with modified shoots)