Objectives Cell Division & Meristems • Chromosomal & non-chromosomal events of cell division • Types of primary meristems • The tunica-corpus concept (shoots) • “Open” & “Closed” meristems (roots) February 1, 2006 • Distinguish between primary & secondary growth • Anatomy of the root surface Mitotic Stages in Onion Root Roles During Plant Cell Division Chromosomes & their replication during cell cycle (M) Mitosis Other Roles During Cell Division • Microtubules – spindle; guiding dictyosome vesicles to cell plate; pre-prophase band. • Microfilaments – guide new wall materials into place on membranes. • Phragmoplast – aggregation of microtubules to cell plate; callose often present. • Cytokinesis – non-nuclear cell division. (G2) Gap 2 (G1) Gap 1 (S) Phase DNA synthesis Cells that cease division Concepts of Meristematic Shoot Organization •Wolff (1759) recognized plant apex as site of primary plant growth. •Hanstein (1870) developed “histogen” theory (dermatogen, periblem, plerome) in roots. •Schmidt (1924) concept of tunica-corpus structure & growth in shoots. 1 Types of Primary Meristems Apical Meristem: A complex of cells composed of initials & their immediate derivatives. • Shoot apical • Root apical • Lateral (shoots & roots) • Intercalary (rib/file) • Calyptrogen Intercalary Meristems • Occur near tip of shoot or root • Cell division is localized • Derivatives also divide before differentiation • Cell division may also occur at distances away from apex in a “wave effect” Pea Meristems Derived from the apical meristem; continuing division from some distance. May be at single site, or multiple ones (waves). Zone of Cell Elongation Zone of Cell Division Primary Meristems (a.k.a. Histogens) •Protoderm: epidermis & rhizodermis •Ground meristem: pith, cortex, endodermis Anticlinal vs. Periclinal Divisions •Procambium: vascular tissues, pericycle •Calyptrogen: rootcap 2 Root & Shoot Meristems Shoot Apical Meristem (SAM) 1. Single Apical Cell – Found in non-seed plants, e.g. Psilotum, Equisetum, & many ferns. Promeristem – Initiating cells & their recent derivatives Primary meristems – Initial differentiation into Protoderm, Procambium & Ground Meristem (& Calyptrogen) Interpretation of Tunica-Corpus Concept 2. Non-stratified – Found in most gymnosperms A. Initial layer B. Central mother cell zone C. Peripheral zone originating leaves D. Rib meristem, forming pith 3. Stratified (Tunica-Corpus) – Found in most angiosperms A. Tunica zone of 1-5 layers (mostly 2) with anticlinal divisions B. Corpus zone of random orientation Gymnosperm Shoot Apex (Non-Stratified) Tunica = Protoderm plus additional cells (Calyptrogen in root) Corpus = Central mother cells, rib meristem, flank meristem Tunica-Corpus Rib (File) Meristem 3 Root Quiescent Centers -- Not Found in Shoots Root Meristem 1. Single Apical Cell – Equisetum & some ferns. Apical cell forms derivatives from each side. • Closed meristem Central cells in distinct layers (stratified) 2. Multiple Initials – (closed) 2 or 3 distinct layers of cells giving rise to vascular cylinder, cortex & root cap. (Found in both monocots & dicots) Open meristem Cells ahead of central cells act as columella mother cells 3. Common Initials (open meristems) – all root tissues from a common group of unstratified initials. (Found in both monocots & dicots) Tobacco Separate initials for central cylinder, cortex & rootcap. (Closed meristem) Lateral cells act as cortical & protoderm initials Corn Root Apical Meristem Onion Common initials. (Open or transversal meristem) Soybean Root: Conventional SEM vs. Cryo Terms to Know • Growth: Increase in size by cell division &/or cell enlargement • Differentiation: Physiological & Conventional morphological change usually associated with specialization • Development: Change in form & complexity from beginning to maturity • Morphogenesis: Combination of Cryo development & differentiation to generate new form 4