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BIOL 1030 TOPIC 4 LECTURE NOTES

Topic 4: Plant Diversity II - Seed Plants (Chs. 30, 38)


I.

Seed plants
A. all are heterosporous vascular plants
B. three major reproductive adaptations
1. gametophyte reduced to dependence on sporophyte; retained in moist reproductive tissue
2. seed baby plant in a lockbox with its lunch; highly resistant structures that allow for a dormant phase in the life
cycle to wait out poor environmental conditions
3. evolution of pollen as male gametophyte many seed plants are no longer tied to external water for fertilization
C. common ancestor with seeds gave rise to all seed plants (gymnosperms and angiosperms); together, gymnosperms and
angiosperms are a monophyletic group
D. fossil evidence indicates origins in progymnosperm group about 360 MYA
E. gametophytes are completely dependent on parent sporophytes for nutrition and are composed of only a few cells
1. male gametophytes develop from microspores
become pollen grains
entire male gametophyte moved to the female as pollen grains
cannot perform photosynthesis, depends on nutrients that came from the parent sporophyte
2. female gametophytes develop from megaspores within ovules
ovule contains female gametophyte surrounded by nucellus (megasporangium)
nucellus is surrounded by 1-2 integuments (cell layers that serve as protective covers)
micropyle opening in integuments (allows sperm to get in)
cannot perform photosynthesis, depends on nutrients from the parent sporophyte
3. means of transporting sperm to egg varies, but typically uses a growing pollen tube that does not require outside water
4. moving pollen to vicinity of ovule called pollination; agents include wind, animals
F. seed develops from ovule
1. seeds are highly resistant structures that allow for a dormant phase in the life cycle to wait out poor environmental
conditions
2. embryo protected by a seed coat, an extra layer of hardened tissue derived from sporophyte tissue in the ovule
(sporophyte tissue from parent, not from embryo)
enhanced protection from drought, cold, heat
some protection from pathogens and predators
external water only needed at germination
initial food supply for germinating plant is enclosed
3. seeds replace spores as means of dispersal; can enhance means of dispersal
G. seeds plants together are a monophyletic group
H. divided into two groups based on whether or not ovule is completely enclosed by sporophyte tissue at time of pollination
1. gymnosperms naked seed
2. angiosperms covered seed covered in Topic 7

II. gymnosperms
A. long thought to be a grade, but molecular data shows that living members may actually form a clade
B. 4 phyla with living members
1. essentially, all seed plants that are not angiosperms
2. all lack flowers and fruits that are found in angiosperms
3. ovule not completely enclosed by sporophyte tissue at time of pollination
4. instead, ovule sits exposed on a scale (a modified leaf)
C. 4 phyla
1. Phylum Coniferophyta (the conifers)
2. Phylum Cycadophyta (the cycads)
3. Phylum Ginkgophyta (Ginkgo)
4. Phylum Gnetophyta (the gnetophytes)
III. Phylum Coniferophyta (the conifers)
A. monophyletic group
B. ~600 living species; worldwide distribution, more common in cold or dry regions
C. pines, spruces, firs, cedars, junipers, hemlocks, yews, larches, cypresses, redwoods
D. nearly all are evergreen
E. many have needle-shaped leaves adapted to dry conditions (resistant to water loss)

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BIOL 1030 TOPIC 4 LECTURE NOTES


1. thick cuticle
2. stomata in pits
F. tallest plant: more than 110 m (Coastal Redwood, Sequoia sempervirens)
G. oldest tree: Methuselah, estimated more than 4600 years old (Bristlecone Pine, Pinus longaeva)
H. sources of timber, paper, resin, cancer drug taxol, etc.
I. soft wood (unlike angiosperm trees, no vessels or fibers in xylem)
J. pines as a representative group
1. over 100 species
2. native to Northern hemisphere
3. typically thick bark (survive fires, drought)
4. secrete resin from leaves and bark
response to wounding
deters fungal and insect attacks
source of turpentine (volatile liquid, organic solvent) and solid rosin
K. pine life cycle
1. pine tree is sporophyte, with sporangia located on cones
2. gametophyte generations reduced; retained within sporangia
male gametophyte is pollen grain (no antheridium)
female gametophyte produces archegonia within ovule
3. heterosporous: separate male and female cones
4. male cones (pollen cones)
clusters of 30-70
usually at tips of lower branches
1-4 cm long; papery scales in spirals or whorls
pair of microsporangia sacs within each scale
microspore mother cells in microsporangia form haploid microspores
each microspore becomes 4-celled pollen grain
pollen grain carried by wind (pair of air sacs provides buoyancy)
mature pollen grains have a Mickey Mouse appearance
one cluster of pollen cones can yield over 1 million pollen grains
5. female cones (ovulate cones)
typically on upper branches of same tree with pollen cones
larger than pollen cones
scales become woody (highly lignified)
pair of ovules develop at base of each scale
megasporangium called nucellus embedded in each ovule
nucellus is nutritive tissue surrounded by thick integument (covering) with hole (micropyle) near one end
one layer of integument later becomes seed coat
single megaspore mother cell in each megasporangium
produces 4 haploid megaspores; 3 break down
surviving megaspore develops over about one year into female gametophyte with sometimes thousands of cells
female gametophyte has 2-6 archegonia, at micropylar end
each archegonium has one large egg (visible without a microscope)
female cones take two or more seasons to mature
6. reproduction
scales of ovulate cone open
pollen lands near micropyle, caught by sticky fluid
evaporating fluid pulls pollen through micropyle into ovule
scales close (female gametophyte not mature)
pollen grain germinates, forming pollen tube that digests through nucellus (takes about 15 months to reach
archegonium)
one of the four pollen grain cells (the generative cell) undergoes mitosis; one of products divides again, making two
sperm cells
mature male gametophyte is germinated pollen grain with pollen tube and sperm
when archegonium is reached, one sperm fertilizes egg

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BIOL 1030 TOPIC 4 LECTURE NOTES

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zygote develops into embryo within seed; usually only one successful zygote per ovule
embryo (new sporophyte, 2N) has rudimentary root, several embryonic leaves (cotyledons)
food source in seed derives from rest of female gametophyte (1N)
scales of cone open and separate; winged seeds disperse
from initial ovulate cone formation to final seed production 3 years or more

IV. Phylum Cycadophyta (the cycads)


A. monophyletic group
B. ~200 living species, tropical and subtropical; many in danger of extinction
C. along with conifers, dominated Mesozoic era (245 MYA 65 MYA)
D. slow-growing; some grow >15 m tall
E. most resemble palm trees, but produce cones (female cones up to 45 kg, or 100 lbs!)
F. have life cycle similar to pines
G. unusual sperm
1. have thousands of flagella arranged in spirals
2. swim within ovule to archegonium
3. largest sperm known
V. Phylum Ginkgophyta (Ginkgo)
A. monophyletic group
B. 1 living species, Ginkgo biloba (also known as the maidenhair tree)
C. exists only in cultivation (no natural native populations); first cultivated in Japan and China
D. deciduous lose leaves
E. flagellated sperm (similar to cycads)
F. dioecious
G. stinky seed coverings (produce butyric and isobutyric acid, making the smell of rancid butter)
H. males often planted on city streets (do not stink like females; resistant to air pollution)
VI. Phylum Gnetophyta (the gnetophytes)
A. apparently a monophyletic group, but could be paraphyletic
B. ~70 living species in 3 genera: Welwitschia, Ephedra, Gnetum
C. some evidence that they form a clade with angiosperms (if so, then gymnosperms are what?)
1. vessels in xylem (common in angiosperms, found only in these gymnosperms)
2. members of Gnetum have broad leaves similar to angiosperm leaves
3. some genetic similarity to angiosperms
D. Welwitschia bizarre plants of southwest African deserts
1. stem is shallow cup that tapers into a taproot
2. two leathery leaves (often split) grow continuously from base
3. conelike reproductive structures at leaf base
4. dioecious separate male and female plants
E. Ephedra common in Mexico and southwestern US, but found on most continents
1. shrubby, stems resemble horsetails (jointed, with tiny scale-like leaves at each node)
2. some species monoecious (male and female parts on same plant), some dioecious
3. drug ephedrine historically extracted mainly from a Chinese species of Ephedra
VII.Phylum Anthophyta flowering plants (antho flower)
A. also known as angiosperms (angeion vessel or enclosure; sperma seed)
B. ovules enclosed within carpel (parent diploid sporophytic tissue) at pollination
1. the vessel is the carpel, which is a modified leaf
2. carpels, especially their enlarged basal portion (the ovary), usually develop into fruit, which is unique to angiosperms
C. about 250,000 known living species (dominant photosynthetic organisms on land)
D. predominant source of human food
E. most widespread and diverse plant phylum
1. range from microscopic to plants with leaves over 6 m long
2. flowers show incredible variety from species to species
3. variety of lifestyles includes parasites (ex.: mistletoe, dodder, beechdrops); mycotrophs (derive nutrients from fungi;
ex.: Indian Pipe, others); epiphytes (ex.: some orchids); carnivorous (ex. pitcher plants, sundews, Venus flytrap)
F. monophyletic group with seeds, refined xylem, double fertilization, and these synapomorphic characteristics:
1. seed contains endosperm

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BIOL 1030 TOPIC 4 LECTURE NOTES


2. presence of flowers (modified stems and leaves)
3. true fruits
G. evolutionary history
1. monophyletic group
2. origin about 140 MYA
H. phylogeny
1. historically divided into two classes, dicots and monocots
recent genetic analysis has shown that the traditional dicots are a paraphyletic group
thus, the old classification scheme is being replaced
2. no conclusive cladogram has been produced for angiosperms
studies are ongoing
most modern cladograms have Amborella and water lilies as a sister group (or groups) to the rest of the
angiosperms
cladogram below from http://tolweb.org/tree?group=Angiosperms&contgroup=Spermatopsida

3.

4.

various class-level groupings have been proposed, the overall naming and formal classification within Phylum
Anthophyta is still in a state of flux
nevertheless, by far most of the living angiosperm species are found within two monophyletic groups, eudicots and
monocots
eudicots
most have embryos have two cotyledons (seed leaves)
leaves have netlike veins
flower part typically in multiples of 4 or 5
groups of vascular tissues in a ring
pollen grains mostly with 3 or more apertures
endosperm mostly used up in mature eudicot seeds
about 175,000 living species; includes nearly all flowering trees and shrubs
about a sixth are annuals (entire growth cycle in one year or less)
monocots
embryos have one cotyledon
leaves have essentially parallel veins
flower part typically in multiples of 3
groups of vascular tissues scattered
pollen grains mostly with one aperture
endosperm typically present in mature monocot seeds
about 65,000 living species; no true wood, few annuals

VIII. Why were (and are) angiosperms successful?


A. 130 MYA two major continental masses
1. Laurasia = North America, Europe, Asia
2. Gondwanaland = South America, Africa, Australia, Antarctica, India, New Zealand)
B. angiosperms first appeared in Gondwanaland, in what was likely a drier interior region
C. advantages of flowering plants
1. transfer of pollen over great distances promotes outcrossing
2. efficient seed dispersal via fruit
3. endosperm gives seedlings a fast start

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4. leaves appropriate for fast growth in hot, dry environment
D. coevolution with insects
1. dominant by ~80 MYA, second half of Cretaceous Period
2. all present angiosperm families represented by that time
3. many insect orders appeared or became more abundant at that time
IX. Flowers
A. modified stems with modified leaves
B. develop as primordium bud at end of stalk called pedicel
C. pedicel widens at tip to form receptacle
D. other flower parts attached to receptacle in four whorls; from outside in:
1. calyx sepals; usually green, leaf-like, and protect immature flower
2. corolla petals; usually colorful, attract pollinators; together with calyx called perianth
3. androecium stamens; male reproductive structures
filament + anther
microspores produced within anther, shed as pollen
4. gynoecium female reproductive structure
center location is most protected
formed from leaf-like structure with ovules along margin
edges fold inwards around ovules, forming carpels
primitive: many separate carpels
advanced: carpels fused (called pistil)
carpel/pistil segments
ovary swollen base with 1 to hundreds of ovules; develops into fruit
stigma tip; sticky and/or feathery to catch pollen
style usually present; separates stigma from ovary
nectaries may be present at base of pistil; secrete sugar, amino acids, and other compounds to attract pollinators
E. know the structures of a flower [Figure 38.2] and their functions

X. typical Angiosperm life cycle


A. female gametophyte
1. single diploid megaspore mother cell in ovule undergoes meiosis while flower develops
2. of 4 haploid megaspores produced, usually 3 break down
3. remaining megaspore expands and replicates and divides until there are 8 haploid nuclei in two groups of 4
4. one nuclei from each group migrates toward center; these are polar nuclei
5. polar nuclei usually fuse to make a diploid nucleus, but may remain separate in either case, they wind up in a single
cell
6. cell walls form around other nuclei, creating the 7-celled, 8-nucleate embryo sac or megagametophyte (female
gametophyte)
7. meanwhile, two layers (integuments) of ovule develop into seed coat with micropyle (small opening)
8. in the megagametophyte, one of the cells closest to the micropyle becomes the egg; the other two there are synergids

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9. the three cells on the other end (the antipodals) eventually break down
B. male gametophyte
1. anthers with patches of tissue that become chambers lined with nutritive cells
2. each patch has many diploid microspore mother cells
3. microspore mother cell undergoes meiosis, making 4 haploid microspores that typically remain grouped in a tetrad
4. each microspore nucleus replicates and divides once (via mitosis) without cytokinesis (meaning they remain as one cell
with two nuclei, a binucleate microspore)
5. usually, tetrad then breaks up
6. two-layered wall develops around each binucleate microspore, now called a pollen grain
outer wall sculptured, appearance usually species-specific, often has chemicals that can react with an appropriate
stigma to stimulate pollen tube formation
apertures in outer wall where pollen tube may grow out; eudicots usually 3; monocots usually 1
C. pollination transfer of pollen to a stigma
1. usually between flowers of separate plants
2. agents include wind, water, gravity, mammals, birds, insects
3. various reward systems for animal agents (pollen, nectar, etc.)
4. evolution of floral characteristics associated with pollination
5. some plants self-pollinate (inbreeding) pollen to same plant
6. pollination followed by fertilization only if chemical signals are right
D. fertilization
1. pollen grain cytoplasm absorbs substances from stigma
2. bulge forms through an aperture in pollen grain; becomes pollen tube
3. pollen tube follows chemical gradient through style to micropyle
chemicals diffuse from embryo sac
micropyle usually reached within a few days (up to a year in some species)
4. pollen grain has two nuclei; one, the generative nucleus, lags behind
5. generative nucleus undergoes mitosis to make two non-flagellated sperm; this may occur in pollen grain or in pollen tube
(male gametophyte now mature)
6. pollen tube enters embryo sac, destroying a synergid
7. double fertilization essentially unique to angiosperms
one sperm unites with egg, forming zygote
other sperm unites with polar nuclei, forming 3N primary endosperm
primary endosperm rapidly undergoes many cycles of mitosis, forming endosperm
endosperm provides nutrients for embryo; in many seeds, it is gone by the time the seed is mature
seed coat hardens
remaining haploid cells degenerate
now have seed with 2N embryo, 3N endosperm, and 2N seed coat (seed coat from parent female tissue)
XI. Seeds
A. embryo quickly forms all systems, then growth arrested (dormancy) mature seed about 10% water, very low metabolic
activity
B. typically, dormancy occurs just after first leaves (cotyledons, or seed leaves) form
C. stored food (in angiosperms, 3N endosperm and/or cotyledons)
D. seed coat tough, relatively impermeable
1. protection from predators, pathogens
2. protection from desiccation, harsh conditions (crucial on land)
3. may allow seed to last hundreds of years
E. dormancy broken only when conditions are right (seed bank in soil)
F. germination = breaking dormancy = resuming metabolic activity, growing out of seed coat; occurs after water penetrates
seed coat to embryo, bringing oxygen
XII.Fruits mature ovaries
A. fleshy pomes (apples), drupes (peaches), true berries (blueberries, peppers), hesperidiums (oranges), pepos (melons,
gourds), aggregate fruits (strawberries, raspberries), multiple fruits (pineapple, fig)
B. dry follicles (milkweed, magnolia), legumes (peas, beans), siliques and silicles (mustards), capsules (irises, lilies, orchids),
caryopses (grasses), nuts (chestnuts, hazelnuts, acorns), achenes (sunflowers), samaras (maples, elms, ashes), schizocarps
(parsleys)
C. dispersal

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1.

2.
3.

4.
XIII.
A.
B.
C.
D.
E.
F.

by wind
wings maples
parachutes dandelions, milkweeds
dust-like seeds orchids
by water coconuts
by vertebrates
fleshy, edible fruits (blue, black, red) seeds often deposited in feces
dry, edible nuts, others squirrels bury and forget about them
dry, inedible hooks to grasp hair, feathers (cockleburs, etc.)
by explosive dehiscence (jewelweed, others)

many angiosperms also have asexual (or vegetative) reproduction


stolons runners long slender stems that grow along soil (ex.: strawberry)
rhizomes underground stems common in grasses; bulbs and tubers are rhizomes specialized for storage (ex.: potato)
suckers roots produce sprouts that grow into new plants (ex.: apple, raspberry, banana)
adventitious leaves numerous plantlets develop from tissue in notches along leaves
apomixes embryos in seeds may be produced asexually
artificial: cuttings (for some species, can get roots to grow with appropriate environment)

XIV. Floral Evolution


A. first flowers
1. numerous spirally arranged sepals, petals, stamens, and carpels
2. petals and sepals similar in color and form
3. all parts free (not fused)
B. parts
1. complete calyx + corolla + androecium + gynoecium
2. incomplete one or more whorls absent
3. perfect has both androecium and gynoecium
4. imperfect missing either androecium or gynoecium
5. complete flowers are always perfect; incomplete flowers can be either perfect or imperfect
C. trends
1. separate floral parts grouped together or fused
connation fusing within a whorl
adnation fusing between whorls (for example, sepals and petals fused together)
2. reduction or loss of floral parts
3. bilateral symmetry instead of radial symmetry
ancestral type: radial symmetry; example: buttercups
derived type: bilateral symmetry; examples: snapdragons, orchids
bilateral symmetry in some cases has arisen independently in different groups
XV. Pollination mechanisms (pollination syndromes)
A. wind passive, primitive (oaks, cottonwoods, birches, grasses)
1. copious amounts of pollen
2. most pollen travels no more than 100 m
3. flowers small, greenish, odorless
4. corollas reduced or absent
5. often grouped in large numbers, may hang down with tassels that wave in wind and shed pollen freely
6. male and female parts often well-separated on plant to reduce chance of self-pollination
7. often flower before leaves grow keeps leaves out of the way
B. animals some cycads and gnetophytes also have this, so symplesiomorphic trait
1. bees most numerous of insect-pollinated plants use bees
find via odor
orient via shape, color, and texture
usually blue or yellow flowers, bee sees in ultraviolet
many have stripes or lines of dots to indicate nectaries (nectar guides)
nectar offered as food for bees (pollen also)
often close association between a bee species and a plant species

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flower only open when bees are active
pollen collecting apparatus specific for particular plant
2. other insects
butterflies flower usually has flat landing platform and long, slender floral tubes for long proboscis
moths flower usually white, yellow, or other pale color, heavily scented, typically need to be found at night
flies flower usually smells and somewhat appears like feces or rotting meat
beetle large flowers, copious pollen; beetle may eat other flower parts
3. birds
large amounts of nectar
red bees cant see red, less likely to feed on the copious nectar
usually odorless birds have a poor sense of smell
often in long, thick tube
4. mammals (bats especially) uncommon, but for some species is the only means of pollination; variety of appearances
C. self-pollination
1. small, inconspicuous flowers
2. shed pollen directly onto stigma (or falls there by gravity); often before bud opens
3. advantageous occasionally because no other plant is needed and no vector is needed good when pollinators arent
around (Artic, mountains)
4. if you are well-adapted, might as well produce clones
5. disadvantage of genetic load of bad mutations
XVI.
A.
B.
C.

Promoting outcrossing
staminate and pistillate flowers
dioecious separate sexes
monoecious
1. dichogamous stamens and pistils reach maturity at different times
2. stigma and stamens dont touch (includes heterostyle)
3. genetic self-incompatibility pollen tube arrested or never germinates

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