Professional Documents
Culture Documents
3Bio01
Histogen Theory
Plant tissue derived from the subapical meristem divided into precursors. 1. Dermatogen- the outermost layer, it gives rise to the protoderm. 2. Plerome- the central cylinder, it gives rise to the procambium. 3. Periblem- sandwiched between the dermatogen and plerome, it gives rise to the ground meristem. 4. Calyptrogen- most distal, it gives rise to the root cap.
Roots can have open or closed organization at the root tip. How a root tip is organized can best be seen with a median longitudinal section.
Closed organization means that the files of cells that arise from the root tip can be traced back to meristematic layers, or histogens. For example, the vascular tissue in this diagram can be clearly traced back to a single layer of cells. This histogen is called the plerome. The ground tissue in this drawing originates from a layer called the periblem, and the epidermis and the root cap both share a histogen layer which is called the calyptrogen/ dermatogen complex or dermatocalyptrogen.
In open organization, the differentiated cell files cannot be traced back to a single, distinguishable layer.
Quiescent Center
The center of the RAM is occupied by a quiescent center which has low mitotic activity. The quiescent center is most apparent in actively growing roots and is lost during dormancy, carbohydrate starvation or root cap removal. Evidence suggests the quiescent center does function as the zone of initials. Infrequent division of initial cells in the quiescent center is the source of cells for the root apical meristem. These initial cells and tissue patterns become established in the embryo in the case of the primary root, and in the new lateral meristems in the case of secondary roots.
its own meristem that pushes cells forward into the cap. As they move through the cap, these cells differentiate into columella cells. Columella cells each contain 15-30 amyloplasts that sediment in response to gravity to the lower side of the cell. Besides protecting the growing root tip and its meristem, the root cap senses light and pressure exerted by soil particles. Within a few days, columella cells differentiate into peripheral cells. The peripheral cells of the root cap and the epidermal cells of the root produce and secrete large amounts of mucigel, a slimy substance made by their dictyosomes. Root cap cells only last 2-3 weeks because as the cells become secretory, the middle lamella weakens, the cells separate and are sloughed off into the soil. Mucigel is a hydrated polysaccharide containing sugars, organic acids, vitamins, enzymes, and amino acids.
Root Development
A small number of stem cells at the tip of the root generate all of the cell types through stereotyped divisions followed by cell differentiation and regulated cell expansion. Because root growth is indeterminate, these processes are continual, resulting in all developmental stages being present at all times. The radial symmetry of the root combined with a lack of cell movement means that clonally related cells are frequently found in cell files. These cell files can be traced back to their origins, which are four types of stem cells (or initial cells) at the root tip. The epidermal/lateral root cap initials give rise to the epidermis and the outer portion of the root cap known as the lateral root cap. The central portion of the root cap, the columella initials. The ground tissue cells, the cortex and endodermis, are generated by division of the cortical/endodermal initials. In older roots, the cortex and endodermis probably have their own committed stem cells.
Finally, the vascular tissue and pericycle have their own central cylinder initials which produces the procambium. The procambium divides to produce the protoxylem, which in turn forms the metaxylem and xylem, and protophloem, which in turn forms the metaphloem and phloem. The metaxylem enlarges and dies after depositing the secondary wall. Internal to and contacting all the initials is a small number of central cells that are mitotically inactive and are known as the quiescent center (QC). Division of initials can be either solely anticlinal (orthogonal to the axis of growth) resulting in a single file of cells or first anticlinal then periclinal (parallel to the axis of growth) resulting in two or more cell layers. The columella initials generally divide only anticlinally and their progeny undergo rapid cell expansion and then differentiate, producing starch-containing amyloplasts that play a role in gravity sensing. The other three types of initials generally undergo both anticlinal and periclinal divisions, resulting in cell lineages that acquire different identities.
Lateral Roots
Lateral roots extend horizontally from the primary root and serve to anchor the plant securely into the soil. This branching of roots also contributes to water uptake, and facilitates the extraction of nutrients required for the growth and development of the plant. Many different factors are involved in the formation of lateral roots. Regulation of root formation is tightly controlled by plant hormones such as auxin, and by the precise control of aspects of the cell cycle. Such control can be particularly useful: increased auxin levels, which help to promote lateral root development, occur after the formation of the leaf primordia which are able to synthesize the hormone. This allows coordination of root development with leaf development, enabling a balance between carbon and nitrogen metabolism to be established.
Like shoot branching, root branching can be either terminal or lateral, with the terminal mode being more common in lower plants and lateral much more common in angiosperms. a. Terminal branching involves the division of the RAM into 2 with the subsequent production of 2 roots. b. Lateral branching is different in roots than in shoots. 1. Lateral roots initiate from internal cells of the pericycle. Initiation occurs in the late cell elongation/early cell differentiation zone, in pericycle cells that are partially to fully differentiated. Thus there is no detached meristem.
2.
3.
4.
A small group of pericycle cells reorient their axis of polarity to the radial dimension and begin growing and dividing to form a mound of cells. With continued growth and division, the mound of cells becomes organized into a RAM with root cap, while still within the tissues of the main root. Continued growth allows the lateral root to penetrate the endodermis, cortex and epidermis, finally reaching the exterior of the parent root.
Initiation of secondary growth takes place in the zone of maturation soon after the cells stop elongating there. The vascular cambium differentiates between the primary xylem and phloem in this zone and pericycle cells divide simultaneously with the procambium initials. The result is a cylinder of cambium encircling the primary xylem. The vascular cambium almost immediately begins producing xylem cells inward and phloem cells toward the outside of the root, in the process flattening the primary phloem against the more resistant endodermis. Concomitant differentiation of cork cambia in the pericycle adds other areas of cell division in the stele.
The combination of periderm and vascular tissue production not only physically breaks the remaining cells of the cortex and epidermis, but the lignified and suberized new cell walls laid down by the cambia effectively isolate the outer tissues as well from their source of supplies in the interior of the root. By the end of the first year, secondary growth has obliterated all but the central core of primary xylem cells and a few fibers of primary xylem pushed against the periderm. The zones at this time, therefore, from outside to inside are: periderm, pericycle, primary phloem, secondary phloem, vascular cambium, secondary xylem, and primary xylem.
References
y http://www-plb.ucdavis.edu/labs/rost/tomato/Roots/roottip.html y http://www.public.iastate.edu/~bot.512/lectures/Roots.htm y http://www2.mcdaniel.edu/Biology/botf99/rootuse/rootanatomy.html y http://en.wikipedia.org/wiki/Meristem#Root_apical_meristems y http://www.bio.mtu.edu/~hlyoungs/plant_development/root_dev.pdf y http://vannocke.hrt.msu.edu/plb865/Root%20structure%20and%20de y y y y