WB01343_.gif (599 bytes)     Coniferophyta     WB01345_.gif (616 bytes)

The Coniferophyta is a relatively largeGymStrobusLarge150.jpg (60841 bytes) group of plants which forms the dominant component of vast ecosystems, especially in the northern hemisphere (Northern Boreal Forest). Some of these plants are the largest organisms on the planet and also have the greatest longevity. The most significant vegetative adaptations of this taxon include Secondary Growth and the production of protective Buds. They also have a significant amount of internodal elongation which allows them to grow faster than the other plants we have reviewed. Their Leaves have a battery of adaptations which foster survival in extreme habitats. The "tree line" in the northern hemisphere is the conifer GymTalLarch200.jpg (84633 bytes)line as these are the most tolerant woody plants with regard to heat, cold and aridity.

Link to Secondary Growth WB01339_.gif (896 bytes)

Secondary Growth is the result of Lateral Meristems (Vascular Cambium & Cork Cambium). The word Cambium signifies the meristematic nature of these two. The plane of cell division in both is predominately Periclinal. This produces radial files of cells which increase the girth of the organ in which they occur.

The Vascular Cambium is a continuation of the Procambium. However, it induces cambial activity inGyMwoodring200Crop.jpg (56241 bytes) adjacent cells and thus adds to its circumference. The Cork Cambium arises de novo (new) from Parenchyma.

The Vascular Cambium produces Secondary Xylem (wood) and Secondary Phloem (inner bark).

Secondary Xylem provides structural support but it also provides a conduit for the translocation of water throughout the entire plant. The largest conifers are several hundred feet high and their root systems are equally large if not of greater extent. The Tracheary ElementsGymSequoiaGrand150.jpg (66813 bytes) in conifers are Tracheids. These serve the dual functions of the secondary xylem but they represent a compromise in terms of both.Tracheid properties differ depending on the environmental conditions under which they are formed. Tracheids have a thicker wall and a narrower diameter if they are formed during periods of water or temperature stress. The opposite occurs during times of plenty. The former are better for support and the latter are better for conduction. Angiosperms produce Vessel Members and Fibers which are more specialized for conduction or support, respectively, and are thus more efficient at both.

Secondary Phloem transports sucrose throughout the organism. Its Sieve Elements are Sieve Cells. These have Sieve Pores in their walls. They are lined withGymBarkMetaSequoia250.jpg (105971 bytes) Callose and they are  similar to sieve cells in seedless vascular plants. Angiosperms have Sieve Tube Members which are more efficient for conduction.

The Cork Cambium (Phellogen) produces Cork (Phellem). Cork is commonly known as "outer bark". This tissue is dead, except for the Phellogen and its immediate derivatives. The walls of Cork Cells are impregnated with Suberin. This is a hydrophobic material that is water-proof and pathogen-proof. It prevents excess water loss and it also insulates living cells from the external environment. Some plants like Douglas Fir and Ponderosa Pine haveGymPonderosaHabitatCrop200.jpg (80225 bytes) barks which protect them from prairie fires. This allows them to survive in environments which are hostile to their competitors.

The Apical Meristems of Conifers are multicellular. They have much in common with the apical meristems of flowering plants. They do not have noticeable "Initials". The Shoot Apical Meristem has a prominent Quiescent Center which has been called the Central Mother Cell Zone. The Root Apical Meristem tends to be broad and all of the tissues of the Root Body and Root Cap are produced by the same Meristem Proper.

GymRootMaeristem200.jpg (83441 bytes)
Typical Conifer Root Apical Meristem
GymSAM300.jpg (84517 bytes)
Typical Conifer Shoot Apical Meristem
GymRAMNom300.jpg (90510 bytes)
Typical Conifer Root Apical Meristem
GymBuds240.jpg (25920 bytes)
Bud Scales enclose Shoot Apices and protect them from environmental trauma.
GymRootLatWholCrop.jpg (40437 bytes)
Root Branching is Lateral rather than Dichotomous.
VegBudExposed200Lac.jpg (36260 bytes)
Each Vegetative Bud contains a legion of preformed leaf primordia. These grow and differentiate rapidly once the bud "flushes" in the spring

(Temp. Zone).
GymTapRoot.jpg (14662 bytes)
Conifers produce a strong tap root system that becomes highly branched and extensive.
GymGrowthFlush200.jpg (46295 bytes)
The enclosed stems of Buds, exhibit explosive growth when permissive conditions return and the buds burst open. Stem branching is Lateral.
GymAraucariaTreesCrop100.jpg (32288 bytes)
Lateral Branching is obvious in Araucaria

(Norfolk Island Pine).
GymSequoiaSAMXSModLab.jpg (208210 bytes)
The Leaf Primordia are tightly clustered around the Shoot Apical Meristem (SAM)
Most Conifers have Needle-like or Scale-like Leaves
GymConesMegaNeedle250Lab.jpg (92649 bytes) GymCypressLeaves300.jpg (56075 bytes)
Needle-like Leaves of Pinus Scale-like Leaves of Cypress

Conifer Leaves have many adaptations that equip them to deal with dry, hot and cold environments.

They have a reduced Lamina (blade).

Pinus monophylla has a cylindrical shape. This give it a small Surface Area/Volume ratio. This means that evaporation will be minimized compared to a flat blade.

If the separate needles of 2-5 needle Pines are viewed together they form a cylinder.

The Epidermis has a thick Cuticle & its walls may be Lignified.

Stomata are "sunken".

A Sclerified Hypodermis (layer(s) below Epidermis) is usually present.

Closely spaced Chlorenchyma

An Endodermis separates the  Mesophyll & Vascular Bundle.

The Vascular Tissues are unbranched and reside in the center of the leaf.

Cross Section of a Pine NeedleCoNeedleXS400Lab.jpg (140058 bytes)
Gym3NeedComp300.jpg (69471 bytes)
Cross Section through the Leaves of a 3-Needle Pine: These form a de facto cylinder.
GYMNeedleX-SPmonticila200Lab.jpg (98417 bytes)
Cross Section of a Pine Needle
GymPiNeedCloseLab400.jpg (202391 bytes)
Part of a Pine Needle showing the Hypodermis and Sunken Stomata.

Stems

Cross section of a young Pinus stem: Note the Interrupted ring of Vascular Tissues GymPinusYunStemXSColrLab400.jpg (224882 bytes)
Pinus stem after a period of Secondary Growth: The Secondary Xylem has stained Red. GymStemX-STwig240Lab.jpg (140406 bytes)
Pinus Stem after two years of growth GymTwigTwoYearPinusLab.jpg (126535 bytes)

All Conifers produce Periderm. Periderm usually arises in the subepidermal Parenchyma. Several cells divide Periclinally and continue to divide this way. This results in radial files of cells which emanate from these Meristematic Cells which are called the Cork Cambium or Phellogen. The Phellogen tends to produce cells towards the outside of the Stem or Root. Meristematic activity spreads laterally until the entire circumference of the organ is encompassed. Cork Cells (Phellem) have Suberized walls and are dead at maturity. Suberin is waxy and prevents excess evaporation. It is almost indigestible by microbes and thus provides a barrier against pathogens. The absence of protoplasm in the dead cork cells deprives pathogens of any nutrients. It also insures that there are no Symplastic connections through which viruses might spread. Commercial Cork comes from the "Cork Oak". It has insulating properties because air is trapped within the individual cork cells. Consequently, the Periderm has insulating properties which lend some protection against temperature extremes in nature. The Periderm is an extremely important adaptation.

PeridermEarlyLab300.jpg (56519 bytes)
Early stage in Periderm (Cork) development: Note the radial arrangement of the Cork cells. The red cells contain Suberin.
GymBarkX-S200Lab.jpg (72394 bytes)
Thick Periderm from Ponderosa Pine: The light lines represent isolated Cork Cambia.

The Primary Root of Conifers is similar to what we have seen before. However, Conifer Roots have Secondary Growth. They have Secondary Vascular Tissues and Periderm. This allows them to expand over time and thus increase their ability to anchor the ever-increasing above ground biomass of the shoot system. It also allows them to persist for centuries in some cases. The Periderm has functions similar to the Periderm on aerial structures. Secondary Vascular Tissues also have the same primary functions as their aerial counterparts.

GymGinkgoRoot240Lab.jpg (62824 bytes)
GymGinkgoRootSecGrow.jpg (293251 bytes)
Cross Section of a Conifer Root with Secondary Growth

WB01342_.gif (412 bytes)