‘OLUTION AND DIVERSITY OF VASCULAR PLANTS

. 2005. Phylogeny of cryptogrannnoid ferns and related taxa

‘tp://amerfernsoc.org
based on rbcL sequences Nordic Journal

lists many resources, including pubiications,

ites, commerjcal fern sites, and fern databases.
of

references, local, national, and interna

I
5
EVOLUTION AND DIVERSITY OF
WOODY AND SEED PLANTS
129 Ginkgophyta 144
LlGNOPHYTESWOODY PLANTS
Ginkgoaceae 145
SPERMATOPHYTES—SEED PLANTS 131 145
Coniferae—Conifers
Seed Evolution 131 148
Pinopsida
Pollination Droplet 135 148
Pinaceae
Pollen Grains 135 151
Cupressopsida
Pollen Tube 136 151
Araucariaceae
Ovule and Seed Development 136 151
Cupressaceae
Seed Adaptations 139 154
Podocarpaceae
Eustele 139 Taxaceae 154
Gnetales 156
DIVERSITY OF WOODYAND SEED PLANTS 139
Ephedraceae 157
Archeopteris 139
“Pteridosperms”—”Seed Ferns” 139 REVIEW QJESTIONS 160
Gymnospermae—Gymnosperms 140 160
EXERCISES
Cycadophyta—Cycads 140
REFERENCES FOR FURTHER STUDY 161
Cycadaceae 141
142 WEB SITES 162
Zamiaceae

LIGNOPHYTIES—WOODY PLANTS
The lignophytes, or woody plants (also called Lignophyta),
are a monophyletic lineage of euphyllous vascular plants that
share the derived features of a vascular cambium, which
gives rise to wood, and a cork cambium, which produces
cork (Figures 5.1, 5.2). Growth of the vascular and cork
cambia is called secondary growth because it initiates after
the vertical extension of stems and roots due to cell expansion
(primary growth). A 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 in extant,
eustelic spermatophytes (see later discussion). The cells of
the vascular cambium divide mostly tangentially (parallel to
a tangential plane), resulting initially in two concentric layers
of cells (Figure 5.3A). One of these layers remains as the vas
cular cambium and continues to divide indefinitely; the other
layer eventually differentiates into either secondary xylem
wood, if produced to the inside of the cambium, or secondary
phloem, if produced to the outside (Figure 5.3A,B). Because
layers of cells are produced both to the inside and outside of
a continuously generated cambium, this type of growth is
termed bifacial. Generally, much more secondary xylem is
produced than secondary phloem. [Note that a secondary
cambium independently evolved in fossil lineages within the
lycophytes (e.g., Lepidodendron) and equisetophytes (e.g.,
Calamites), but this cambium was unifacial, producing sec
ondary xylem (wood) to the inside but no outer secondary
phloem, likely limiting in terms of an adaptive feature.]
Secondary growth results in an increase of the width or girth
of stems and roots (Figures 5.3B, 5.4). This 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).
A cork cambium is similar to a vascular cambium, only it
differentiates near the periphery of the stem or root axis. The
cork cambium and its derivatives constitute the periderm
(referred to as the outer bark). The outermost layer of the
periderm is cork (Figure 5.3B). Cork cells contain a waxy

129
© 2010 Elsevier Inc. All rights reserved.
doi: 10.101 61B978-0- 12-374380-000005.2
 ______ __
____I

UNIT II EVOLUTION AND DIVERSITY OF PLANTS 131

130 CHAPTER 5 EVOLUTION AND DIVERSITY OF WOODY AND SEED PLANTS

- Lignophyta (Woody Plants) classification and identification of woody plants. Wood ana
tomical features may also be used to study the past, a spe
Spermatophyta (Seed Plants) cialty known as dendrochronolOgy (see Chapter 10).
Gymnospermae (Gymnosperms) Another feature of lignophytes is that they possess ances
shoot
trally monopodial growth, in which a single main
Coniferae (Conifers) (see
develops branches from lateral (usually axillary) buds
monopodial growth is presumed to
Chapters 4, 9). Although
t Cupressopsida monilophyte—lignophyte split, it
have arisen prior to the
a) o a) S of forming
a a
C r Gnetales —,
a)
S
o o
— enabled woody plants in particular the capability
extensive (sometimes massive) woody branching systems,
a) 0 0 a)
C: S a)
a) t
permitting them to survive and reproduce more effectively.
I
4., 0-I. a) :t S a)
Ct S rj
C C S S Ct
o 0
a)
is 4.
a) a)
C — •0
oO
a)
.sç a)
a) S
C
-5c .

— (/,
C)
- 0
(/D
5
‘1 SPERMATOPHYTESSEED PLANTS —

epimatium
+ F
The spermatophyta, commonly called spermatophytes
seed plants, are a monophyletic lineage within the lignophytes
or

receptacle -
this
(Figure 5.1). The major evolutionary novelty that unites
is an
iiiiifiiiii 1 ovule/scale group is the seed. A seed is defined as an embryo, which
porose sur
I
1g., -,
/
immature diploid sporophyte developing from the zygote,
-

enveloped by a seed coat

rounded by nutritive tissue and
W

immature
(Figure 5.5). The embryo generally consists of an
called the
root called the radicle, a shoot apical meristem
pollen tube—sperm nonmotile (siphonogamy) cotyledons;
epicotyl, and one or more young seed leaves, the
iiiiiiiifiiii leaves simple V c •4L
%H
the transition region between root and stem
hypocotyl (Figures 5.5, 5.10). An immature seed,
is called the
prior to

I fertilization, is known as an ovule.

-

:14
I’

— s eustele
SEED EVOI.,UTION
The
The evolution of the seed involved several steps.
— — pollen tube—sperm motile (zooidogamy) or more
exact sequence of these is not certain, and two
— — endosporic, male gametophyte = pollen grain occurred concomitantly
“steps” in seed evolution may have
in seed
— — pollination droplet and be functionally correlated. The probable steps
— — integument with micropyle evolution are as follows (Figure 5.6):
— — retention of megaspore within megasporangium
1. HeterospOry. Heterospory is the formation of two types
— — reduction to 1 megaspore per megasporangium SEED large,
(embryo of haploid spores within two types of sporangia:
— — endosporic female gametophyte via meiosis
a,, fewer-numbered megaspores, which develop
-

+ nutntive tissue •‘
— — heterospory pa-. ,41 t numerous
+ integuments)
lIst. in the megasporangium, and small, more
t =extincttaxon
— — cork cambium (periderm) !%: r microsporeS, the products of meiosis in the microsporafl
— which a
gium (Figures 5.6, 5.7). The ancestral condition, in
= extinct lineage vascular cambium (secondary vascular tissue, md. wood)
Each
single spore type forms, is called “homosporyY
.— -a -

-‘- . -

megaspore develops into a female gametophyte that bears

-

FIGURE 5.1 Cladogram of the woody and seed plants. Major apomorphies are indicated beside a thick hash mark. Families in bold are
game
described in detail. Modified from Bowe et al. (2000); Chaw et al. (2000); Frohlich et al. (2000); and Samigullin et al. (1999). only archegonia; a microspore develops into a male
Although heterospory
tophyte, bearing only antheridia.

polymer called suberin (similar to cutin) that is quite resist-
ant to water loss (see Chapter 10).
The vascular cambium and cork cambium constituted major
evolutionary novelties. Secondary xylem, or wood, functions
in structural support, enabling the plant to grow tall and acquire
massive systems of lateral branches. Thus, the vascular cam-
bium was a precursor to the formation of intricately branched
Et :rLct
plants
shrubs or trees with tall overstory canopies (e.g., Figure 5.2), was prerequisite to seed evolution, there are fossil
among
a significant ecological adaptation. Cork produced by the cork that were heterosporous but had not evolved seeds,
(Figures 5.1, 5.1 3A;
cambium functions as a thick layer of cells that protects these being species of Archeopteris
has evolved
the delicate vascular cambium and secondary phloem from see later discussion). Note that heterospory
independently in other, nonseed plants, e.g., in the
extant
mechanical damage, predation, and desiccation. ferns
Wood anatomy can be quite complex. The details of
FIGURE 5.2 Composite
massive, nonclonal
lycophytes Selaginella and Isoetes and in the water
giant sequoia, a woody conifer that is the most
cellular structure are important characters used in the (Chapter 4).
organism on Earth, and among the tallest of trees.
AND DIVERSITY OF WOODY AND SEED PLANTS UNIT II EVOLUTION AND DIVERSITY OF PLANTS 131

— Lignophyta (Woody Plants) — —

classification and identification of woody plants. Wood ana
1
Spermatophyta (Seed Plants) tomical features may also be used to study the past, a spe
cialty known as dendrochronology (see Chapter 10).
- Gymnospermae (Gymnosperms) —

Another feature of lignophytes is that they possess ances

1 trally monopodial growth, in which a single main shoot
— Conjferae (Conifers) —
—

develops branches from lateral (usually axillary) buds (see

1 — Cupressopsida Chapters 4, 9). Although monopodial growth is presumed to
have arisen prior to the monilophyte—lignophyte split, it
r Gnetales -, a:
a.)
a:
a.)
C.)
a.)
a.)
a.)
a.) a:
—
enabled woody plants in particular the capability of forming
C.)
a: a:
a a: extensive (sometimes massive) woody branching systems,
a: a: a: permitting them to survive and reproduce more effectively.
.
C.? C C
— -a:
C.) a:
a.,) c_)
SPERMATOPHYTES—SEED PLANTS
aril
The Spermatophyta, commonly called spermatophytes or
seed plants, are a monophyletic lineage within the lignophytes
(Figure 5.1). The major evolutionary novelty that unites this

S group is the seed. A seed is defined as an embryo, which is an

immature diploid sporophyte developing from the zygote, sur
rounded by nutritive tissue and enveloped by a seed coat
(Figure 5.5). The embryo generally consists of an immature
pollen tube—sperm nonmotjie (siphonogamy) root called the radicle, a shoot apical meristem called the
epicotyl, and one or more young seed leaves, the cotyledons;
the transition region between root and stem is called the
hypocotyl (Figures 5.5, 5.10). An immature seed, prior to
fertilization, is known as an ovule.

eustele SEED EVOLUTION

I
— —
—
pollen tube—sperm motile (zooidogamy) The evolution of the seed involved several steps. The
endosporic, male gametophyte pollen grain exact sequence of these is not certain, and two or more
pollination droplet “steps” in seed evolution may have occurred concomitantly
integument with micropyle and be functionally correlated. The probable steps in seed
retention of megaspore within megasporangjum evolution are as follows (Figure 5.6):
reduction to 1 megaspore per megasporangjum SEED 1. Heterospory. Heterospory is the formation of two types
endosporic female gametophyte (embryo of haploid spores within two types of sporangia: large,
+ nutritive tissue
heterospory
+ integuments)
fewer-numbered megaspores, which develop via meiosis
cork cambjum (periderm) in the megasporangium, and small, more numerous
vascular cambjum (secondary vascular tissue, mci. wood) microspores, the products of meiosis in the microsporan
glum (Figures 5.6, 5.7). The ancestral condition, in which a
plants. Major apomorphies are indicated beside a single spore type forms, is called “homospory.” Each
thick hash mark. Families in bold are
10); Chaw et al. (2000); Frohljch et al. (2000); megaspore develops into a female gametophyte that bears
and Samigullin et al. (1999).
only archegonia; a microspore develops into a male game
tophyte, bearing only antheridia. Although heterospory
is quite resist shrubs or trees with tall overstory canopies (e.g., Figure 5.2),
was prerequisite to seed evolution, there are fossil plants
a significant ecological adaptation. Cork produced by the
cork that were heterosporous but had not evolved seeds, among
flstituted major cambium functions as a thick layer of cells that protects
these being species of Archeopteris (Figures 5.1, 5.13A;
‘ood, functions the delicate vascular cambium and secondary phloem from
tall and acquire see later discussion). Note that heterospory has evolved
mechanical damage, predation, and desiccation. independently in other, nonseed plants, e.g., in the extant
vascular cam Wood anatomy can be quite complex. The details of FIGURI 5.2 Composite photograph ofSequoiadendrongiganteum,
lycophytes Selaginella and Isoetes and in the water ferns
ately branched cellular structure are important characters used in giant sequoia, a woody conifer that is the most massive, nonclonal
the (Chapter 4).
organism on Earth, and among the tallest of trees.
 NTS 133
UNIT II EVOLUTION AND DIVERSITY OF PLA
132 CHAPTER 5 EVOLUTION AND DIVERSITY OF WOODY AND SEED PLANTS

I FIGURE 5.4

2. Endospory. Endospory is
Woody stem cross-section, Pinus. sp. A. One

the complete development of,

year’s growth. B. Four years’ growth.

increased availability of space and

megasporangium.
resources in the

hyte within the original

in this case, the female gametop ad of the megaspore
condition, in which 4. Retention of the megaspore. Inste
spore wall (Figure 5.6). The ancestral ancestral condi
an external gameto being released from the sporangium (the
the spore germinates and grows as eed plants),
ution of endosporic tion, as occurs in all homosporous nons
phyte, is called exospory. The evol megasporangium
with that of in seed plants it is retained within the
female gametophytes was correlated by a reduction in
grains); see later (Figure 5.6). This was accompanied
endosporic male gametophytes (pollen
thickness of the megaspore wall.
discussion. opyle. The final
ber to one. Reduction of 5. Evolution of the integument & micr
3. Reduction of megaspore num event in seed evolution was the enve
lopment of the
. First, the number
megaspore number occurred in two ways e, calle d the integu
undergo meiosis megasporangium by a layer of tissu
of cells within the megasporangium that s from the base of
aspore mother ment (Figure 5.6). The integument grow
(each termed a megasporocyte or meg called a nucellus
(Figure 5.6). This the megasporangium (which is often
cell) was reduced, from several to one
A single diploid megasporocyte gives rise
to four haploid when surrounded by an integument) and
sugg
envelopes it,
ests that the
haploid meg aspores pro except at the distal end. Fossil evidence
megaspores. Second, of the four lobes derived
t, leavi ng only integument likely evolved from separate
duced by meiosis, three consistently abor surrounded the
megaspore also from telomes (ancestral branches) that
vascular one functional megaspore. This single ovules prior to
lated with the megasporangium. These “preovules”, i.e.,
cambjum
undergoes a great increase in size, corre a rim or ring of
2 xylem the evolution of integuments, possessed
the lagenos
tissue at the apex of the megasporangium,
n grains to a pol
tome, which functioned to funnel polle
art and Rothwell 1993
lination chamber. (See, e.g., Stew
occurred with
2 phloem
for details.) The epitome of seed evolution
es to form the
the evolutionary “fusion” of the telom
completely sur
integument, a continuous sheath that
nt of all extant seed
rounds the nucellus. The integume
radicle
periderrn called the micro-
plants has a small pore at the distal end
stral lagenostome
embryo
pyle. The micropyle replaced the ance
in angiosperms, of
as the site of entry of pollen grains (or
also functions in the
fluthtive tissue pollen tubes). The micropyle
ation and resorp
(female gametophyle cotyledons mechanics of pollination droplet form
B cork
or endosperm)
tion (see below). Note that a single integ
ument represents
s; in angiosperms
(epidermis sloughed off to outside) the ancestral condition of spermatophyte
later (Chapter 6).
trated here. a second integument layer evolved
FIGURE 5.3 A. Development of the vascular cambium. B. Development of secondary vascular tissue in the stem, illustrated here for a FIGURE 5.5 Morphology of a seed. Pinus sp. illus
eusteljc stem.
AND DIVERSITY OF WOODYAND SEED PLANTS UNIT II EVOLUTION AND DIVERSITY OF PLANTS 133

1
FIGURE 5.4 Woody stem cross-section, Pinus. sp. A. One year’s growth. B. Four years’ growth.

2. Endospory. Endospory is the complete development of, increased availability of space and resources in the
in this case, the female gametophyte within the original megasporangium.
spore wall (Figure 5.6). The ancestral condition, in which 4. Retention of the megaspore. Instead of the megaspore
the spore germinates and grows as an external gameto being released from the sporangium (the ancestral condi
phyte, is called exospory. The evolution of endosporic tion, as occurs in all homosporous nonseed plants),
female gametophytes was correlated with that of in seed plants it is retained within the megasporangium
endosporic male gametophytes (pollen grains); see later (Figure 5.6). This was accompanied by a reduction in
discussion. thickness of the megaspore wall.
3. Reduction of megaspore number to one. Reduction of 5. Evolution of the integument & micropyle. The final
2’ xylem megaspore number occurred in two ways. First, the number event in seed evolution was the envelopment of the
vascular vascular of cells within the megasporangium that undergo meiosis megasporangium by a layer of tissue, called the integu
cambium cambium 2 phloem (each termed a megasporocyte or megaspore mother ment (Figure 5.6). The integument grows from the base of
cell) was reduced, from several to one (Figure 5.6). This the megasporangium (which is often called a nucellus
single diploid megasporocyte gives rise to four haploid when surrounded by an integument) and envelopes it,
megaspores. Second, of the four haploid megaspores pro except at the distal end. Fossil evidence suggests that the
duced by meiosis, three consistently abort, leaving only integument likely evolved from separate lobes derived
l’phloem one functional megaspore. This single megaspore also from telomes (ancestral branches) that surrounded the
undergoes a great increase in size, correlated with the megasporangium. These “preovules”, i.e., ovules prior to
2’ xylem
1’ xylem the evolution of integuments, possessed a rim or ring of
tissue at the apex of the megasporangium, the lagenos
tome, which functioned to funnel pollen grains to a pol
seed coat
lination chamber. (See, e.g., Stewart and Rothwell 1993
2’ phloem
for details.) The epitome of seed evolution occurred with
the evolutionary “fusion” of the telomes to form the
integument, a continuous sheath that completely sur
radicle
periderm rounds the nucellus. The integument of all extant seed
plants has a small pore at the distal end called the micro-
cortex embryo pyle. The micropyle replaced the ancestral lagenostome
epicotyl as the site of entry of pollen grains (or in angiosperms, of
nutritive tissue
(female gametophyte pollen tubes). The micropyle also functions in the
or endosperm) cotyledons mechanics of pollination droplet formation and resorp
cork tion (see below). Note that a single integument represents
(epidermis sloughed off to outside)
the ancestral condition of spermatophytes; in angiosperms
ambium. B. Development of second vascular tissue in
the stem, illustrated here for a FIGURE 5.5 Morphology of a seed. Pinus sp. illustrated here. a second integument layer evolved later (Chapter 6).
 134 CHAPTER 5 EVOLUTION AND DIVERSITY OF WOODY AND SEED PLANTS UNIT II EVOLUTION AND DIVERSITY OF PLANTS 135

Sporophyte Body
,.._-‘ (2n)
antheridia mitosis, growth, & differentiation mitosis, growth, & differentiation

7
Embryo
(2n) Microsporangium Megasporangium
(2n) (2n)
/ \ \

/
mitosis, growth, & differentiation mitosis, growth, & d(fferentianon
male gametophyte

(n)
‘\ Megasporocyte
SPOROPHYTE GENERATION Microsporocyte
Zygote (2n)
(2n) (2N) (2n)
microsporangium

——fertilization
t rneiosis —— meioszs ——

gametophyte

(n) SpOraflgium
/‘
‘jN
)
Egg
(n)
I)

1.
GAMETOPHYTE GENERATION

Sperm (sperm nonflagellate in

(N)

(n) j Conifers (mci. Gnetales) © (

Microspores
(n) (n)
Jn
Megaspores..)

©
k
1/
and Angiosperms)
female gametophyte
(lost in the Angiosperms
& some Gnetales)
{ Archegonium Antheridium 1 (reduced to absent in
(n) (n) 3’
extant seed plants) mitosis, growth, & differentiation

1. Heterospory megasporangium mitosis, growth, & derentiaiion

Male Gametophyte
(n)

archegonia Female Gametophyte

archegonia (n)

/ FIGURE 5.7 Life cycle of heterosporous seed plants.

0
wall

female gametophyte POLLINATION DROPLET POLLEN GRAINS

(contained in megaspore)
One possible evolutionary novelty associated with seed evo
megasporangiurn megasporangium lution is the pollination droplet. This is a droplet of liquid
that is secreted by the young ovule through the micropyle
2. Endospory 3. Reduction to 1 megaspore (Figures 5.1OA, 5.171). This droplet is mostly water plus
some sugars or amino acids and is formed by the breakdown
of cells at the distal end of the megasporangium (nucellus).
The cavity formed by this breakdown of cells is called the
pollination chamber (Figure 5. bA). The pollination drop
let functions in transporting pollen grains through the micro-
pyle. This occurs by resorption of the droplet, which “pulls”
megaspore pollen grains that have contacted the droplet into the pollina
gametophyte tion chamber. It is unknown whether a pollination droplet
was present in the earliest seed plants. However, the presence
of a pollination droplet in many nonflowering seed plants
megasporangium
suggests that its occurrence may be apomorphic for at least
the extant seed plant lineages. Note that the ovules of
angiosperms lack pollination droplets or pollination cham
bers, as flowering plants have evolved a different mechanism
4. Retention of megaspore 5. Evolution of Integument & Micropyle of pollen grain transfer (see Chapter 6).
FIGURE 5.6 Ovule and seed evolution in the spermatophytes (hypothetical, for purpose of illustration).
Concomitant with the evolution of the seed was the evolution of
pollen grains (Figure 5.8). A pollen grain is, technically, an
immature, endosporic male gametophyte. Endospory in pollen
grain evolution was similar to the same process in seed evolu
tion, involving the development of the male gametophyte within
the original spore wall. Pollen grains of seed plants are extremely
reduced male gametophytes, consisting of only a few cells.
They are termed “immature” male gametophytes because, at the
time of their release, they have not fully differentiated.
After being released from the microsporangium, pollen must
be transported to the micropyle of the ovule (or, in angio
sperms, to the stigmatic tissue of the carpel; see Chapter 6) in
order to ultimately effect fertilization. Wind dispersal, in com
bination with an ovule pollination droplet (see later discus
sion), was probably the ancestral means of pollen transport.
After being transported to the ovule (or stigmatic tissue), the
male gametophyte completes development by undergoing
additional mitotic divisions and differentiation. The male
gametophyte grows an exosporic pollen tube, which functions
 134 CHAPTER 5 EVOLUTION AND DIVERSITY OF WOODY AND SEED PLANTS UNIT II EVOLUTION AND DIVERSITY OF PLANTS 135

Sporophyte Body
,.._-‘ (2n)
antheridia mitosis, growth, & differentiation mitosis, growth, & differentiation

7
Embryo
(2n) Microsporangium Megasporangium
(2n) (2n)
/ \ \

/
mitosis, growth, & differentiation mitosis, growth, & d(fferentianon
male gametophyte

(n)
‘\ Megasporocyte
SPOROPHYTE GENERATION Microsporocyte
Zygote (2n)
(2n) (2N) (2n)
microsporangium

——fertilization
t rneiosis —— meioszs ——

gametophyte

(n) SpOraflgium
/‘
‘jN
)
Egg
(n)
I)

1.
GAMETOPHYTE GENERATION

Sperm (sperm nonflagellate in

(N)

(n) j Conifers (mci. Gnetales) © (

Microspores
(n) (n)
Jn
Megaspores..)

©
k
1/
and Angiosperms)
female gametophyte
(lost in the Angiosperms
& some Gnetales)
{ Archegonium Antheridium 1 (reduced to absent in
(n) (n) 3’
extant seed plants) mitosis, growth, & differentiation

1. Heterospory megasporangium mitosis, growth, & derentiaiion

Male Gametophyte
(n)

archegonia Female Gametophyte

archegonia (n)

/ FIGURE 5.7 Life cycle of heterosporous seed plants.

0
wall

female gametophyte POLLINATION DROPLET POLLEN GRAINS

(contained in megaspore)
One possible evolutionary novelty associated with seed evo
megasporangiurn megasporangium lution is the pollination droplet. This is a droplet of liquid
that is secreted by the young ovule through the micropyle
2. Endospory 3. Reduction to 1 megaspore (Figures 5.1OA, 5.171). This droplet is mostly water plus
some sugars or amino acids and is formed by the breakdown
of cells at the distal end of the megasporangium (nucellus).
The cavity formed by this breakdown of cells is called the
pollination chamber (Figure 5. bA). The pollination drop
let functions in transporting pollen grains through the micro-
pyle. This occurs by resorption of the droplet, which “pulls”
megaspore pollen grains that have contacted the droplet into the pollina
gametophyte tion chamber. It is unknown whether a pollination droplet
was present in the earliest seed plants. However, the presence
of a pollination droplet in many nonflowering seed plants
megasporangium
suggests that its occurrence may be apomorphic for at least
the extant seed plant lineages. Note that the ovules of
angiosperms lack pollination droplets or pollination cham
bers, as flowering plants have evolved a different mechanism
4. Retention of megaspore 5. Evolution of Integument & Micropyle of pollen grain transfer (see Chapter 6).
FIGURE 5.6 Ovule and seed evolution in the spermatophytes (hypothetical, for purpose of illustration).
Concomitant with the evolution of the seed was the evolution of
pollen grains (Figure 5.8). A pollen grain is, technically, an
immature, endosporic male gametophyte. Endospory in pollen
grain evolution was similar to the same process in seed evolu
tion, involving the development of the male gametophyte within
the original spore wall. Pollen grains of seed plants are extremely
reduced male gametophytes, consisting of only a few cells.
They are termed “immature” male gametophytes because, at the
time of their release, they have not fully differentiated.
After being released from the microsporangium, pollen must
be transported to the micropyle of the ovule (or, in angio
sperms, to the stigmatic tissue of the carpel; see Chapter 6) in
order to ultimately effect fertilization. Wind dispersal, in com
bination with an ovule pollination droplet (see later discus
sion), was probably the ancestral means of pollen transport.
After being transported to the ovule (or stigmatic tissue), the
male gametophyte completes development by undergoing
additional mitotic divisions and differentiation. The male
gametophyte grows an exosporic pollen tube, which functions
 136 CHAPTER 5 EVOLUTION AND DIVERSITY OF WOODY AND SEED PLANTS UNIT II EVOLUTION AND DIVERSITY OF PLANTS 137

micropyle

integument (2n)
pollen
pollen
grains
archegonial chamber
pollination droplet
pollination archegonium
- integument chamber (with egg)

S
A ICL mitosiS and

differentiation
(2n)

FIGURE 5.8 Pollen grains—immature male gametophytes of seed plants. A. Zamia sp., a cycad. B. Ginkgo biloba. C. Pinus sp., a conifer. megasporangium female
(nucellus) functional
megaspore gametophyte
(2n) (n)
(n)
as a haustorial organ, obtaining nutrition by absorption from the marriage). A type of siphonogamy evolved independently in
surrounding sporophytic tissue (Figure 5.9; see Pollen Tabe). the angiosperms. In angiosperms, however, the pollen tubes
grow through stylar tissue prior to delivering the sperm to the
A
POLLEN TUBE micropyle
egg of a female gametophyte (see Chapter 6).
The male gametophytes of all extant seed plants form a pollen
tube (Figure 5.9) soon after the pollen grains make contact OVULE AND SEED DEVELOPMENT seed coat’
with the megasporangial (nucellar) tissue of the ovule. In After pollination, the megasporocyte develops within the radicle
extant seed plants the ancestral type of pollen type (found in megasporangium of the ovule (Figures 5.1OA, 5.11A). The mitosis and
cycads and ginkgophytes) was haustorial, in which the male megasporocyte is a single cell that undergoes meiosis, producing egg) hypocotyl
gametophyte feeds (like a parasite) off the tissues of the a tetrad of four haploid megaspores, which in most extant seed megaSporangiUm embryo
nucellus. Motile sperm is delivered from this male gameto plants are arranged in a straight line, or linearly (Figure 5.IOA). (nucellus) (new 2n)
phyte into a fertilization chamber, where the sperm swims to (2n)
The three megaspores that are distal (away from the ovule base) epicotyl
the archegonium containing the egg, a process known as abort; only the proximal megaspore (near the ovule base) con (shoot apex)
zooidogamy (zoom, animal + gamos, marriage). In the coni tinues to develop. In the pollination chamber, the resorbed cotyledons
female
fers (including Gnetales), pollen tubes are also haustorial, but gametophyte
pollen grains (Figures 5. iDA, 5.1 1A) develop into mature male (n) female
deliver nonmotile sperm cells to the archegonium or egg, a gametophytes and form pollen tubes, which grow into the tissue gametophyte
(n)
process known as siphonogamy (siphono, tube + gamos, of the megasporangium (Figures 5. iDA, 5.1 1B). In gymnosperms
B megaspOrangiUm
(degenerate)

FIGURE 5.10 A. Ovule development in the nonflowering spermatophytes. B. Seed development.

micropyle megasporangium
these male gametophytes may live in the megasporangial tissue
for some time, generally several months to a year.
The functional megaspore greatly expands, accompanied
pollen tube
(haustorial) by numerous mitotic divisions, to form the endosporic
female gametophyte (Figures 5.1OA, 5.11B,C). In the seeds
of gymnosperms, archegonia differentiate at the apex of the
germination & female gametophyte (Figure 5.11C,D). As in the nonseed
land plants, each archegonium has a large egg cell and a
differentiation sperm short line of neck cells (plus typically a ventral canal cell or
nucleus). Eventually, the male gametophytes either release
motile
pollen grain mature male sperm cell motile sperm cells (in cycads and Ginkgo) into a cavity
(immature endosporic
male gametophyte)
gametophytes, each
with pollen tube between the megasporangium and female gametophyte
(known as the archegonial chamber; Figure 5.1 DA), or the
FIGURE 5.9 Male gametophyte morphology and development in the nonflowering spermatophytes; Cycas sp., illustrated. (Reproduced
pollen tube of the male gametophyte delivers sperm cells
and modified from Swamy, B. G. L. 1948. American Journal of Botany 35: 77—88, by permission.)
directly into the archegonial neck (in conifers). (Note that
the ovules of some Gnetales and all angiosperms lack arche
gonia.) The end result is that a sperm cell from the male
gametophyte fertilizes the egg of the female gametophyte. A
long period of time (perhaps a year or more) may ensue
between pollination, which is delivery of the pollen grains
to the ovu)e, and fertilization, actual union of sperm and
egg. Note: This is not true for the flowering plants, in which
fertilization generally occurs very soon after pollination (see
Chapter 6).
The resulting diploid zygote, once formed, undergoes
considerable mitotic divisions and differentiation, eventually
maturing into the embryo, the immature sporophyte (Figures
5.1DB, ShE). The tissue of the female gametophyte contin
ues to surround the embryo (Figure 5.11E) and serves as
nutritive tissue for the embryo upon seed germination (except

1
 136 CHAPTER 5 EVOLUTION AND DIVERSITY OF WOODY AND SEED PLANTS UNIT II EVOLUTION AND DIVERSITY OF PLANTS 137

micropyle

integument (2n)
pollen
pollen
grains
archegonial chamber
pollination droplet
pollination archegonium
- integument chamber (with egg)

S
A ICL mitosiS and

differentiation
(2n)

FIGURE 5.8 Pollen grains—immature male gametophytes of seed plants. A. Zamia sp., a cycad. B. Ginkgo biloba. C. Pinus sp., a conifer. megasporangium female
(nucellus) functional
megaspore gametophyte
(2n) (n)
(n)
as a haustorial organ, obtaining nutrition by absorption from the marriage). A type of siphonogamy evolved independently in
surrounding sporophytic tissue (Figure 5.9; see Pollen Tabe). the angiosperms. In angiosperms, however, the pollen tubes
grow through stylar tissue prior to delivering the sperm to the
A
POLLEN TUBE micropyle
egg of a female gametophyte (see Chapter 6).
The male gametophytes of all extant seed plants form a pollen
tube (Figure 5.9) soon after the pollen grains make contact OVULE AND SEED DEVELOPMENT seed coat’
with the megasporangial (nucellar) tissue of the ovule. In After pollination, the megasporocyte develops within the radicle
extant seed plants the ancestral type of pollen type (found in megasporangium of the ovule (Figures 5.1OA, 5.11A). The mitosis and
cycads and ginkgophytes) was haustorial, in which the male megasporocyte is a single cell that undergoes meiosis, producing egg) hypocotyl
gametophyte feeds (like a parasite) off the tissues of the a tetrad of four haploid megaspores, which in most extant seed megaSporangiUm embryo
nucellus. Motile sperm is delivered from this male gameto plants are arranged in a straight line, or linearly (Figure 5.IOA). (nucellus) (new 2n)
phyte into a fertilization chamber, where the sperm swims to (2n)
The three megaspores that are distal (away from the ovule base) epicotyl
the archegonium containing the egg, a process known as abort; only the proximal megaspore (near the ovule base) con (shoot apex)
zooidogamy (zoom, animal + gamos, marriage). In the coni tinues to develop. In the pollination chamber, the resorbed cotyledons
female
fers (including Gnetales), pollen tubes are also haustorial, but gametophyte
pollen grains (Figures 5. iDA, 5.1 1A) develop into mature male (n) female
deliver nonmotile sperm cells to the archegonium or egg, a gametophytes and form pollen tubes, which grow into the tissue gametophyte
(n)
process known as siphonogamy (siphono, tube + gamos, of the megasporangium (Figures 5. iDA, 5.1 1B). In gymnosperms
B megaspOrangiUm
(degenerate)

FIGURE 5.10 A. Ovule development in the nonflowering spermatophytes. B. Seed development.

micropyle megasporangium
these male gametophytes may live in the megasporangial tissue
for some time, generally several months to a year.
The functional megaspore greatly expands, accompanied
pollen tube
(haustorial) by numerous mitotic divisions, to form the endosporic
female gametophyte (Figures 5.1OA, 5.11B,C). In the seeds
of gymnosperms, archegonia differentiate at the apex of the
germination & female gametophyte (Figure 5.11C,D). As in the nonseed
land plants, each archegonium has a large egg cell and a
differentiation sperm short line of neck cells (plus typically a ventral canal cell or
nucleus). Eventually, the male gametophytes either release
motile
pollen grain mature male sperm cell motile sperm cells (in cycads and Ginkgo) into a cavity
(immature endosporic
male gametophyte)
gametophytes, each
with pollen tube between the megasporangium and female gametophyte
(known as the archegonial chamber; Figure 5.1 DA), or the
FIGURE 5.9 Male gametophyte morphology and development in the nonflowering spermatophytes; Cycas sp., illustrated. (Reproduced
pollen tube of the male gametophyte delivers sperm cells
and modified from Swamy, B. G. L. 1948. American Journal of Botany 35: 77—88, by permission.)
directly into the archegonial neck (in conifers). (Note that
the ovules of some Gnetales and all angiosperms lack arche
gonia.) The end result is that a sperm cell from the male
gametophyte fertilizes the egg of the female gametophyte. A
long period of time (perhaps a year or more) may ensue
between pollination, which is delivery of the pollen grains
to the ovu)e, and fertilization, actual union of sperm and
egg. Note: This is not true for the flowering plants, in which
fertilization generally occurs very soon after pollination (see
Chapter 6).
The resulting diploid zygote, once formed, undergoes
considerable mitotic divisions and differentiation, eventually
maturing into the embryo, the immature sporophyte (Figures
5.1DB, ShE). The tissue of the female gametophyte contin
ues to surround the embryo (Figure 5.11E) and serves as
nutritive tissue for the embryo upon seed germination (except

1
138 CHAPTER 5 EVOLUTiON AND DIVERSITY OF WOODYAND SEED PLANTS UNIT II EVOLUTION AND DIVERSITY OF PLANTS 139

integument integument rchegonia

I female
gametophyte gametophyte
1, phloem

1xylern

4
I’, /

V
.
cortex

7
a
pith
female
gametophyte
vascular
A B C bundle

outside. B. Helianthus stem

FIGURE 5.12 Eustele. A. Diagram of eustele. Note single ring of vascular bundles, with xylem inside, phloem
associated fibers.
cross-section, an example of a eustele. C. Close-up of vascular bundle, showing xylem, phloem, and

/--
megasporocyte
in the flowering plants; see Chapter 6). The megasporangium EUSTELE
In addition to the seed, an apomorphy for spermatophytes is

female
11 (nucellus) eventually degenerates. The integument matures
into a peripheral seed coat, which may differentiate into
various hard andlor fleshy layers.
the eustele (Figure 5.12). A eustele is a primary stem vascu
lature (“primary” meaning prior to any secondary growth)
that consists of a single ring of discrete vascular bundles.
r.

gametophyte embryo
Each vascular bundle contains an internal strand of xylem
/ SEED ADAPTATIONS
The adaptive significance of the seed is unquestioned. First,
seeds provide protection, mostly by means of the seed coat,
and an external strand of phloem that are radially oriented,
i.e., positioned along a radius (Figure 5.12).
U-. from mechanical damage, desiccation, and often predation. The protoxylem of the vascular bundles of a eustele is
endarch in position, i.e., toward the center of the stem. This is
- Second, seeds function as the dispersal unit of sexual repro
duction. In many plants the seed has become specially modi distinct from the exarch protoxylem of the lycophytes and the
fied for dispersal. For example, a fleshy outer seed coat layer mesarch protoxylem of most monilophytes (Chapter 4) and of
nuêleus
may function to aid in animal dispersal. In fact, in some plants some fossil relatives that diverged prior to the seed plants.
the seeds are eaten by animals, the outer fleshy layer is
digested, and the remainder of the seed (including the embryo
protected by an inner, hard seed coat layer) passes harmlessly DIVERSITY OF WOODY
through the gut of the animal, ready to germinate with a built AND SEED PLANTS
in supply of fertilizer. In other plants, differentiation of the
seed coat into one or more wings functions in seed dispersal ARCHEOPTEPJS
-1 by wind. Third, the seed coat may function in dormancy A well-known lignophyte that lacked seeds was the fossil plant
Archeopteris (not to be confused with the very famous fossil,
-

••• ‘
‘\sIrile mechanisms that ensure germination of the seed only under
•
.
‘.:;. reptilian bird Archeopteryx). Archeopteris was a large tree, with
ideal conditions of temperature, sunlight, or moisture. Fourth,
...

cells
upon germination, the nutritive tissue surrounding the embryo wood like a conifer but leaves like a fern (Figure 5.13A,B).
provides energy for the young seedling, aiding in successful Sporangia, producing spores, were born on fertile branch
E establishment. systems. Some species of Archeopteris were heterosporous.

FIGURE 5.11 Ovule and seed development, illustrated by Pinus sp. A. Young ovule, longitudinal-section, at time of pollination. Pollen
Interestingly, in seed plants the female gametophyte (which
develops within the megaspore) remains attached to and “PTERIDOSPERMS”—”SEED FERNS”
grains are pulled into micropyle by resorption of pollination droplet. Meiosis of the megasporocyte has yet to occur. B. Post-pollination,
nutritionally dependent upon the sporophyte. This is exactly The “pteridosperms,” or “seed ferns:’ are almost certainly
showing development of the female gametophyte and haustorial pollen tube growth of the male gametophytes within tissue of megasporangium
(nucellus). C. Mature ovule, showing two functional archegonia within female gametophyte. D. Close-up of archegonia, each containing the reverse condition as is found in the liverworts, homworts, a paraphyletic group of fossil plants that had femlike foliage, yet
a large egg cell with a surrounding layer of sterile cells and apical neck. E. Seed longitudinal-section, seed coat removed, showing embryo and mosses (Chapter 3). bore seeds. Medullosa is a well-known example of a seed fern
and surrounding nutritive layer of female gametophytic tissue.
138 CHAPTER 5 EVOLUTiON AND DIVERSITY OF WOODYAND SEED PLANTS UNIT II EVOLUTION AND DIVERSITY OF PLANTS 139

integument integument rchegonia

I female
gametophyte gametophyte
1, phloem

1xylern

4
I’, /

V
.
cortex

7
a
pith
female
gametophyte
vascular
A B C bundle

outside. B. Helianthus stem

FIGURE 5.12 Eustele. A. Diagram of eustele. Note single ring of vascular bundles, with xylem inside, phloem
associated fibers.
cross-section, an example of a eustele. C. Close-up of vascular bundle, showing xylem, phloem, and

/--
megasporocyte
in the flowering plants; see Chapter 6). The megasporangium EUSTELE
In addition to the seed, an apomorphy for spermatophytes is

female
11 (nucellus) eventually degenerates. The integument matures
into a peripheral seed coat, which may differentiate into
various hard andlor fleshy layers.
the eustele (Figure 5.12). A eustele is a primary stem vascu
lature (“primary” meaning prior to any secondary growth)
that consists of a single ring of discrete vascular bundles.
r.

gametophyte embryo
Each vascular bundle contains an internal strand of xylem
/ SEED ADAPTATIONS
The adaptive significance of the seed is unquestioned. First,
seeds provide protection, mostly by means of the seed coat,
and an external strand of phloem that are radially oriented,
i.e., positioned along a radius (Figure 5.12).
U-. from mechanical damage, desiccation, and often predation. The protoxylem of the vascular bundles of a eustele is
endarch in position, i.e., toward the center of the stem. This is
- Second, seeds function as the dispersal unit of sexual repro
duction. In many plants the seed has become specially modi distinct from the exarch protoxylem of the lycophytes and the
fied for dispersal. For example, a fleshy outer seed coat layer mesarch protoxylem of most monilophytes (Chapter 4) and of
nuêleus
may function to aid in animal dispersal. In fact, in some plants some fossil relatives that diverged prior to the seed plants.
the seeds are eaten by animals, the outer fleshy layer is
digested, and the remainder of the seed (including the embryo
protected by an inner, hard seed coat layer) passes harmlessly DIVERSITY OF WOODY
through the gut of the animal, ready to germinate with a built AND SEED PLANTS
in supply of fertilizer. In other plants, differentiation of the
seed coat into one or more wings functions in seed dispersal ARCHEOPTEPJS
-1 by wind. Third, the seed coat may function in dormancy A well-known lignophyte that lacked seeds was the fossil plant
Archeopteris (not to be confused with the very famous fossil,
-

••• ‘
‘\sIrile mechanisms that ensure germination of the seed only under
•
.
‘.:;. reptilian bird Archeopteryx). Archeopteris was a large tree, with
ideal conditions of temperature, sunlight, or moisture. Fourth,
...

cells
upon germination, the nutritive tissue surrounding the embryo wood like a conifer but leaves like a fern (Figure 5.13A,B).
provides energy for the young seedling, aiding in successful Sporangia, producing spores, were born on fertile branch
E establishment. systems. Some species of Archeopteris were heterosporous.

FIGURE 5.11 Ovule and seed development, illustrated by Pinus sp. A. Young ovule, longitudinal-section, at time of pollination. Pollen
Interestingly, in seed plants the female gametophyte (which
develops within the megaspore) remains attached to and “PTERIDOSPERMS”—”SEED FERNS”
grains are pulled into micropyle by resorption of pollination droplet. Meiosis of the megasporocyte has yet to occur. B. Post-pollination,
nutritionally dependent upon the sporophyte. This is exactly The “pteridosperms,” or “seed ferns:’ are almost certainly
showing development of the female gametophyte and haustorial pollen tube growth of the male gametophytes within tissue of megasporangium
(nucellus). C. Mature ovule, showing two functional archegonia within female gametophyte. D. Close-up of archegonia, each containing the reverse condition as is found in the liverworts, homworts, a paraphyletic group of fossil plants that had femlike foliage, yet
a large egg cell with a surrounding layer of sterile cells and apical neck. E. Seed longitudinal-section, seed coat removed, showing embryo and mosses (Chapter 3). bore seeds. Medullosa is a well-known example of a seed fern
and surrounding nutritive layer of female gametophytic tissue.