Professional Documents
Culture Documents
Lignophytes and Spermatophytes (Chap 5)
Lignophytes and Spermatophytes (Chap 5)
E VOLUTION AND D IVERSITY OF W OODY AND SEED P LANTS (C HAPTER 5) P AGES 129-162
F OR A MORE ELABORATE DISCUSSION AND ILLUSTRATIONS , PLEASE REFER TO YOUR TEXTBOOK.
. (PAGE 129)
Monophyletic lineage of euphyllous vascular plants sharing derived features: o Vascular Cambium gives rise to wood o Cork Cambium produces cork (Figures 5.1, 5.2 page 130, 131) Secondary Growth is what we call the growth of the vascular and cork cambium o it is because it initiates after the vertical extension of stems and roots due to cell expansion (primary growth) y Vascular Cambium is a sheath, or hollow cylinder, of cells that develops within the stems and roots as a continuous layer, between the xylem and phloem. o Cells of the vascular cambium divide mostly tangentially (parallel to a tangential plane), resulting initially in two concentric layers of cells (Figure 5.3A page 132) One of these layers remains as the vascular cambium and continues to divide indefinitely The other layer eventually differentiates into either: y Secondary Xylem if produced to the inside of the cambium y Secondary Phloem if produced to the outside (Figure 5.3A,B) Bificial a type of growth wherein layers of cells are produced both to the inside and outside of a continuously generated cambium. *Note: more 2o xylem is produced than 2o phloem
y o 2o cambium independently evolved in fossil lineage w/in the lycophytes, but this cambium was unificial, producing 2o xylem but not 2o phloem
Secondary growth results in an increase of the width or girth of stems and roots Occurs both by expansion of the new cells generated by the cambium and by accompanying radial divisions, increasing the number of cells within a given growth ring. Many woody plants have regular growth periods, e.g., forming annual rings of wood
(Figure 5.4 page 133)
Cork Cambium similar to vascular cambium, only it differentiates near the periphery of the stem or root axis.
Cork cambium and its derivatives constitute the periderm (referred to as the outer bark) Cork the outermost layer of the periderm (Figure 5.3B page 132) y Cork cells contain a waxy polymer called suberin (similar to cutin) that is quite resistant to water loss Secondary xylem, or wood functions in structural support. o
Cork functions as a thick layer of cells that protects the delicate vascular cambium and secondary phloem from mechanical damage, predation, and desiccation. Dendochronology study of the past using the features of the wood anatomy Monopodial Growth another feature lignophytes possess, in which a single main shoot branches from lateral (usually axillary) buds. o It enabled woody plants in particular the capability of forming extensive (sometimes massive) woody branching systems permitting them to survive and reproduce more effectively.
. (PAGE 131)
o Major evolutionary novelty: seed (Figure 5.5 page 133) o Seed an embryo an immature diploid sporophyte developing from the zygote, surrounded by nutritive tissue and enveloped by a seed coat Radicle an immature root Epicotyl a shoot apical meristem Cotyledons one or more young seed leaves Hypocotyl the transition region between root and stem (Figure 5.5, 5.10)
SEED EVOLUTION
y Probable steps in seed evolution are as follows (Figure 5.6 page 134) 1. Heterospory the formation of two types of haploid spores within two types of sporangia: large, fewer-numbered megaspores, which develop via meiosis in the megasporangium, and small, more numerous microspores, the product of meiosis in the microsporangium (Figures 5.6, 5.7 page 135,
135)
Homospory ancestral condition where only a single spore type forms Megaspore develops into a female gametophyte, bearing archegonia Microspore develops into a male gametophyte, bearing antheridia 2. Endospory is the complete development of, in this case, the female gametophyte within the original spore wall (Figure 5.6 page 134) Exospory ancestral condition where spore germinates and grows as an external gametophyte 3. Reduction of megaspore number to one. Occurred in two ways: First: the number of cells within the megasporangium that undergo meiosis (each termed a megasporocyte or megaspore mother cell) was reduced, from several to one. This single diploid megasporocyte gives rise to four haploid megaspores Second: of the four haploid megaspores produced by meiosis, three consistently abort, leaving only one functional megaspore. This single megaspore also undergoes a great increase in size, correlated with the increased availability of space and resources in the megasporangium. 4. Retention of the megaspore. Instead of the megaspore being released from the sporangium (the ancestral condition), in seed plants it is retained within the megasporangium. This was accompanied by a reduction in thickness of the megaspore wall.
5. Evolution of the integument & micropyle. The final event in seed evolution was the envelopment of the megasporangium by a layer of tissue, called the integument. The integument grows from the base of the megasporangium (which is often called a nucellus when surrounded by an integument) and envelopes it, except at the distal end. y Fossil evidence suggest that the integument likely evolved from telomes (ancestral branches) that surrounded the megasporangium y These preovules , i.e., ovules prior to the evolution of integuments possessed a rim or ring of tissue at the apex of the megasporangium, the lagenostome, which functioned to funnel pollen grains to a pollination chamber y Epitome of seed evolution = fusion of the telomes to form the integument a continuous sheath that completely surrounds the nucellus. y Micropyle a small pore at the distal end in the integument of all extant seed plants. functions as the site of entry of pollen grains (or in angiosperms, of pollen tubes). also functions in mechanics of pollination droplet formation and resorption.
which functions as an haustorial organ, obtaining nutrition by absorption from the surrounding sporophytic tissue
o The megasporocyte is a single cell that undergoes meiosis, producing a tetrad of four haploid megaspores, which in most extant seed plants are arranged in a straight line, or linearly (Figure 5.10A
page 137)
o The three megaspores that are distal (away from ovule base) abort; only the proximal megaspore (near the ovule base) continues to develop. o In the pollination chamber, the resorbed pollen grains (Figures 5.10A, 5.11A) develop into mature male gametophytes and form pollen tubes, which grow into the tissue of the megasporangium (Figures
5.10A, 5.11B).
o The functional megaspore greatly expands, accompanied by mitotic divisions, to form the endosporic female gametophyte (Figure 5.10A, 5.11B,C) o In the seeds of gymnosperms, archegonia differentiate at the apex of the female gametophyte
(Figure 5.11C,D)
o As in large egg cell and a short line of neck cells (plus typically a ventral canal cell or nucleus). Eventually, the male gametophytes either release motile sperm cells (in cycads and Ginkgo) into a cavity between the megasporangium and female gametophyte (known as the archegonial chamber; Figure 5.10A), or the pollen tube of the male gametophyte delivers sperm cells directly into the archegonial neck (in conifers). o The end result is that a sperm cell from the male gametophyte fertilizes the egg of the female gametophyte. o A long period of time may ensue between pollination and fertilization (Note: this is not true for the flowering
plants, in which fertilization occurs soon after pollination)
Pollination delivery of the pollen grains to the ovule Fertilization actual union of sperm and egg
o The resulting diploid zygote, once formed, undergoes mitotic divisions and differentiation, eventually maturing into the embryo, the immature sporophyte (Figures 5.10B, 5.11E) o The tissue of the female gametophyte continues to surround the embryo (Figure 5.11E) and serves as nutritive tissue for the embryo upon seed germination (except in flowering plants) o The megasporangium (nucellus) eventually degenerates. o The integument matures into a peripheral seed coat, which may differentiate into various hard and/or fleshy layers.
y y