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Seed Plants
Chapter 26
1
The Evolution of Seed Plants
• Seed plants first appeared 305–465 mya• Success attributed to evolution of seed
– Seed includes embryo, food for the embryo, and a seed coat
– Allows the “clock to be stopped” to survive harsh periods before germinating
– “Naked” seeds first, then later development of fruits enhanced dispersal
2
• Seed plants produce 2 kinds of gametophytes• Male gametophytes
– Pollen grains– Dispersed by wind or a pollinator– No need for water
• Female gametophytes – Develop within an ovule– Enclosed within diploid sporophyte tissue in
angiosperms
3
Five Phyla of Extant Seed Plants
• Coniferophyta• Cycadophyta• Gnetophyta• Ginkophyta• Anthophyta
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5
Gymnosperms
• Plants with “naked seeds”• There are four living groups
• Coniferophytes• Cycadophytes• Gnetophytes• Ginkgophytes
• All lack flowers and fruits of angiosperms• All have the ovule exposed on a “scale”
6
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.
Ang
iosp
erm
s
Gym
nosp
erm
s
Fern
s an
d A
llies
Conifers (phylum Coniferophyta)
• Most diverse and familiar gymnosperm phylum
• Pines, spruces, firs, cedars, and others• Found in colder and sometimes drier
regions of the world• Conifers are sources of important products
– Timber, paper, resin, and taxol (anti-cancer)
7
• Pines– More than 100 species, all
in the Northern hemisphere
– Produce tough needlelike leaves in clusters
– Leaves have thick cuticle and recessed stomata to retard water loss
– Leaves have canals with resin to deter insect and fungal attacks
8
• Pine reproduction• Male gametophytes (pollen grains)
– Develop from microspores in male cones by meiosis
• Female pine cones form on the upper branches of the same tree– Female cones are larger, and have woody
scales– Two ovules develop on each scale– Each contains a megasporangium
• Each will become a female gametophyte
9
10
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.
Microsporemother cell
Microspores PollenAir bladder
Pollination
Pollen tube
SpermMITOSIS
Zygote
Section of seed (second year),showing embryo embeddedin megagametophyte
MITOSIS
Pineseed
MITOSIS
Seedling
Sporophyte
Pollen-bearingcone
Scale
Megasporemother cell
Ovulate(seed-bearing)cone
Megaspore
Pollen tube
n2n
(15 monthsafter pollination)
MEIO
SIS
FERTILIZATION
• Female cones usually take 2 or more seasons to mature• During the first spring, pollen grains drift down between open
scales– Pollen grains drawn down into micropyle– Scales close
• A year later, female gametophyte matures– Pollen tube is digesting its way through– Mature male gametophyte has 2 sperm
• 15 months after pollination, pollen tube reaches archegonium and discharges contents– One sperm unites with egg = zygote– Other sperm degenerates
11
Cycads (phylum Cycadophyta)
• Slow-growing gymnosperms of tropical and subtropical regions
• Sporophytes resemble palm trees
• Female cones can weigh 45 kg
• Have largest sperm cells of all organisms!
12
a.© Luca Invernizzi Tetto/agefotostock
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.
Gnetophytes (phylum Gnetophyta)
• Only gymnosperms with vessels in their xylem
• Contain three (unusual) genera– Welwitschia– Ephedra– Gnetum
13
b.© Juan Carlos Muñoz/agefotostock
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.
Ginkgophytes (phylum Ginkgophyta)
• Only one living species remains– Ginkgo biloba
• Flagellated sperm• Dioecious
– Male and female reproductive structures form on different trees
14
c. © Robert Gustafson/Visuals Unlimited
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.
c.c.
Angiosperms
• Flowering plants • Ovules contain eggs,
and become seeds after egg fertilization
• Carpel, a modified leaf that covers seeds, develops into fruit
15
Ferns and Allies
Angiosperms
Gymnosperms
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.
16
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.
(bottom right): © Goodshoot/Alamy RF
Fusion ofleaf margins
Modified leafwith ovules
Cross section
Ovules(seeds)
Carpel(fruit)
Folding of leafprotects ovules
Ovules
Angiosperm origins are a mystery
– Origins as early as 145–208 MYA
– Oldest known angiosperm in the fossil record is Archaefructus
17
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.
Fruits
© David Dilcher and Ge Sun
Pairedstamens
18
Archaefructus may be the sister clade to all other angiosperms
Gymnosperms AngiospermsGinkgo Gnetophytes Conifers Cycads Archaefructus
(extinct)Amborella Water lillies MonocotsMagnoliidsEudicotsStaranis
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Flowers contain the gametophyte generation
• Flower morphology– All flower parts have evolved either from
stems or leaves– Flower parts are organized in circles called
whorls
19
• Flower whorls– Outermost whorl – sepals– Second whorl – petals– Third whorl – stamens (androecium)
• Each stamen has an anther (where the pollen is) and a filament (stalk)
• Pollen is the male gametophyte– Fourth whorl - gynoecium
• Consists of one or more carpels• Contain the female gametophyte
20
• Carpel has 3 major regions– Ovary – contains the ovules
• The ovary later develops into a fruit, and the ovules inside become the seeds
– Stigma – tip where pollen lands– Style – neck or stalk
21
Carpel
Stigma
Style
Ovule
Ovary wall
StamenAntherFilament
PetalSepal
Ovary
a.
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.
Megaspore (n)
GenerativecellTubenucleus
Pollen tubeSpermTubenucleus
Style
Pollen (n)
Formation ofpollen tube (n)
Polarnuclei
Sperm
Egg
Polarnuclei
Egg
MIT
OSI
S
MIT
OSI
S
Zygote
Embryo (2n)
Cotyledons
Seed (2n)
Endosperm (3n)
Endosperm
Anther
Ovary
Stigma Anther (2n)
Ovule
2n
DO
UB
LEFE
RTI
LIZA
TIO
N
MIT
OSIS
GERMINATIO
N
Microsporemother cells (2n)
Megasporemother cell (2n)
Adult sporophytewith flower (2n)
Youngsporophyte (2n)
Seedcoat
MITOSIS
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22
• Pollination – Mechanical transfer of pollen from anther to
stigma– May or may not be followed by fertilization– Pollen grains develop a pollen tube that is
guided to the embryo sac– The “generative cell” divides to produce two
sperm cells• No flagella on sperm
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• Double fertilization – One sperm unites with egg to form the diploid
zygote (which becomes the new sporophyte embryo)
– Other sperm unites with the two polar nuclei to form the triploid endosperm
• Provides nutrients to new sporophyte embryo
• Seed may remain dormant for many years– Germinate (grow by mitosis) when conditions
are favorable
24
Seeds
• In many angiosperms, development of the embryo stops soon after meristems and cotyledons differentiate
• Integuments develop into a relatively impermeable seed coat
• Encloses the seed with its dormant embryo and stored food
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26
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.
Shoot apical meristem
Root apicalmeristem
Root cap
Endosperm
Procambium
Seed coat(integument)
Cotyledons
• Seeds are an important adaptation1. They maintain dormancy under unfavorable
conditions2. They protect the young plant when it is most
vulnerable3. They provide food for the embryo until it can
produce its own food4. They facilitate dispersal of the embryo
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• Once a seed coat forms, most of the embryo’s metabolic activities cease
• Germination cannot take place until water and oxygen reach the embryo
• Seeds of some plants have been known to remain viable for thousands of years
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• Specific adaptations ensure that seeds will germinate only under appropriate conditions– Some seeds lie within tough cones that do not
open until exposed to fire
29a. b.
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.
a: © Ed Reschke; b: © David Sieren/Visuals Unlimited
Fruits
• Most are simply defined as “mature ovaries” (carpels)
• At the same time seeds are forming, the flower ovary begins to develop into a “fruit”
• However, it is also possible for fruits to develop without seed development– Bananas are propagated asexually
• Not all fruits are something you would want to eat!
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True Berries
The entire pericarp isfleshy, although theremay be a thin skin.Berries have multipleseeds in either one ormore ovaries. The tomato flower had four carpels that fused.Each carpel containsmultiple ovules thatdevelop into seeds.
Outer pericarpSeedFused
carpels
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Drupes
Seed
PericarpExocarp (skin)MesocarpEndocarp (pit)
Single seedenclosed in a hard pit;peaches, plums,cherries. Eachlayer of the pericarp hasa different structureand function, withthe endocarpforming the pit
Samaras
PericarpNot split andwith a wingformed from theouter tissues;maples, elms,ashes.
Seed
(left): © Kingsley Stern; (right): Courtesy of Robert A. Schisling
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.
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Aggregate Fruits
Sepals of asingle flower
Ovary
Seed
Derived from manyovaries of a singleflower; strawberries,blackberries.Unlike tomato,these ovariesare not fusedand coveredby a continuouspericarp.
Multiple Fruits
Individual flowers form fruitsaround a single stem. The fruitsfuse as seen with pineapple. Pericarp of
individual flower
Main stem
(left): Courtesy of Robert A. Schisling; (right): © Charles D. Winters/Photo Researchers, Inc.
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.
Fruit Dispersal
• Occurs through a wide array of methods– Ingestion and transportation by birds or other
vertebrates– Hitching a ride with hooked spines on birds
and mammals– Burial in caches by herbivores– Blowing in the wind– Floating and drifting on water
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