Botany

Chapter 5
Evolution and Diversity of Woody and Seed plants

LIGNOPHYTES—WOODY PLANTS

 Lignophyta – a MONOphyletic lineage of euphyllous

vascular plants
o secondary growth – growth of the vascular and cork
cambia
 initiates after the vertical extension of stems and
roots due to cell expansion (primary growth)
 vascular cambium – shared the derived feature
 gives rise to wood and a cork cambium  Secondary growth – results in an increase of the width or
 cork cambium – produces cork girth of stems and root
o expansion of the new cells generated by the cambium
o radial divisions – increasing the number of cells
within a given growth ring
 regular growth periods – annual rings of wood
 Dendrochronology – tree-ring dating

 cork cambium – similar to a vascular cambium, only it

differentiates near the periphery of the stem or root axis
o periderm – outer bark = cork cambium + cork
o cork – outermost layer of periderm
o suberin – waxy polymer similar to cutin that is quite
resistant to water loss

 Secondary xylem (wood) – functions in structural support,

enabling the plant to grow tall and acquire massive
systems of lateral branches
o vascular cambium – precursor to the formation of
intricately branched shrubs or trees with tall
overstory canopies
o Cork – functions as a thick layer of cells that protects
the delicate vascular cambium and secondary phloem
from mechanical damage, predation, and desiccation

 monopodial growth – a single main shoot develops

branches from lateral (usually axillary) buds
o arisen prior to the monilophyte–lignophyte split
 vascular cambium – a sheath, or hollow cylinder, of cells o enabled woody plants the capability of forming
that develops within the stems and roots as a continuous extensive (sometimes massive) woody branching
layer, between the xylem and phloem in extant, eustelic systems, permitting them to survive and reproduce
spermatophytes more effectively
o cells - divide mostly tangentially (parallel to a
tangential plane), resulting initially in two concentric
SPERMATOPHYTES—SEED PLANTS
layers of cells
 1st layer – remain as the vascular cambium
 Spermatophyta – MONOphyletic
 2nd layer – divide indefinitely – bifacial growth
 secondary xylem = wood – produced to the
inside of the cambium  Seed – major evolutionary novelty
 secondary phloem – produced to the o Embryo – immature diploid sporophyte developing
outside from the zygote, surrounded by nutritive tissue and
o bifacial growth – layers of cells are produced both to enveloped by a seed coat
the inside and outside of a continuously generated  Radicle – immature root
cambium  epicotyl – shoot apical meristem
 more secondary xylem is produced than  cotyledons – young seed leaves
secondary phloem  hypocotyl – transition region b/n root and stem
o ovule – immature seed, prior to fertilization
 Secondary cambium – independently evolved in fossil
lineages within the lycophytes (e.g., Lepidodendron) and
equisetophytes (e.g., Calamites)
o Unifacial growth – produces secondary xylem (wood)
to the inside but no outer secondary phloem

 Seed Evolution
o (1) Heterospory - formation of two types of haploid
spores within two types of sporangia
 (a) megaspores – large, fewer-numbered
 develop via meiosis in the megasporangium
 female gametophyte – bears only
archegonia
 (b) microspores – small, more numerous
 products of meiosis in the
microsporangium
 male gametophyte – bears only antheridia
 homospory – ancestral condition, in which a
single spore type forms
 heterosporous fossil plants w/o seeds
 species of Archeopteris
 heterospory – evolved independently in other,
nonseed plants
 extant lycophytes Selaginella and Isoetes
and in the water ferns
o (2) Endospory – complete development of the female
gametophyte within the original spore wall
 exospory - ancestral condition, in which the spore
germinates and grows as an external gametophyte
 correlated with evolution of endosporic male
gametophytes
o (3) Reduction of megaspore number to one
 megasporocyte (megaspore mother cell) (2n) –
cells within the megasporangium that undergo
meiosis
 reduced from several to one
 gives rise to four haploid megaspores
 four haploid megaspores – three consistently
abort, leaving only one functional megaspore
o (4) Retention of the megaspore
 Megaspore
 released from the sporangium – ancestral
condition
 retained within the megasporangium
 reduction in thickness of the megaspore
wall
o (5) Evolution of the integument & micropyle
 Integument – layer of tissue that envelopes the
megasporangium
 grows from the base of the
megasporangium and envelopes it, except
at the distal end
 a continuous sheath that completely
surrounds the nucellus
 nucellus – megasporangium surrounded by
an integument
 telomes – ancestral branches that surrounded
the megasporangium
 “preovules” – ovules prior to the evolution of
integuments
 Lagenostome – a rim or ring of tissue at the
apex of the megasporangium which
functioned to funnel pollen grains to a
pollination chamber
 Micropyle – a small pore at the distal end of the
integument
 replaced the ancestral lagenostome as the
site of entry of pollen grains (or in
angiosperms, of pollen tubes)
 functions in the mechanics of pollination
droplet formation and resorption
 single integument represents the ancestral
condition of spermatophytes
 in angiosperms. 2nd integument layer
evolved later

POLLINATION DROPLET
 pollination droplet – droplet of liquid that is secreted by
the young ovule through the micropyle
o mostly water + some sugars / amino acids – formed
by the breakdown of cells at the distal end of the
megasporangium (nucellus)
o transports pollen grains through the micropyle
o may be apomorphic for at least the extant seed plant
lineages
 pollination chamber – cavity formed by the breakdown of
cells
 ovules of angiosperms lack pollination droplets or
pollination chambers, as flowering plants have evolved a
different mechanism of pollen grain transfer
POLLEN GRAINS
 pollen grains – an immature, endosporic male
gametophyte
o Endospory in pollen grain evolution – similar to the
same process in seed evolution, involving the
development of the male gametophyte within the
original spore wall
 Pollen grains – extremely reduced male gametophytes,
consisting of only a few cells
o “immature” male gametophytes – not fully
differentiated at the time of their release
 Wind dispersal – ancestral means of pollen transport
 Pollen tube – exosporic which functions as a haustorial
organ, obtaining nutrition by absorption from the
surrounding sporophytic tissue
Pollen – microsporangium → micropyle of the ovule (angiosperms: stigmatic
tissue of the carpel) → male gametophyte – mitotic divisions and
differentiation → pollen tube → sperm → fertilization
POLLEN TUBE
 Haustorial – ancestral type of pollen type in extant seed
plants found in cycads and ginkgophytes
o male gametophyte feeds (like a parasite) off the
tissues of the nucellus
 zooidogamy (zooin, animal + gamos, marriage) – motile
sperm is delivered from this male gametophyte into a
fertilization chamber, where the sperm swims to the
archegonium containing the egg
 siphonogamy (siphono, tube + gamos, marriage)
o in the conifers, including Gnetales
o pollen tubes deliver nonmotile sperm cells to the
archegonium or egg
o A type of siphonogamy evolved independently in the
angiosperms
 pollen tubes grow through stylar tissue prior to
delivering the sperm to the egg of a female
gametophyte
OVULE AND SEED DEVELOPMENT
 megasporocyte – develops within the megasporangium of
the ovule after pollination
o a single cell that undergoes meiosis, producing a
tetrad of four haploid megaspores
 three megaspores that are distal (away from the
ovule base) abort
 proximal megaspore (near the ovule base)
continues to develop
o In the pollination chamber, resorbed pollen grains →
mature male gametophytes → pollen tubes
 In gymnosperms, these male gametophytes may
live in the megasporangial tissue for some time,
generally several months to a year
  functional megaspore – greatly expands, accompanied by
numerous mitotic divisions, to form the endosporic female
gametophyte
o In the seeds of gymnosperms, archegonia
differentiate at the apex of the female gametophyte
o in the nonseed land plants, each archegonium has a
large egg cell and a short line of neck cells (plus
typically a ventral canal cell or nucleus)
 archegonial chamber – cavity between the
megasporangium and female gametophyte
o (in cycads and Ginkgo) male gametophytes release
motile sperm cells
o (in conifers) pollen tube of the male gametophyte
delivers sperm cells directly into the archegonial neck
o **ovules of some Gnetales and all angiosperms lack
archegonia
 Pollination – delivery of the pollen grains to the ovule
 Fertilization – actual union of sperm and egg
o (in flowering plants) fertilization generally occurs very
soon after pollination
Pollen – microsporangium → micropyle of the ovule (angiosperms: stigmatic tissue of
the carpel) → male gametophyte – mitotic divisions and differentiation → pollen tube →
sperm → archegonia → egg cell → fertilization → Zygote (2n) → embryo (immature
sporophyte) → tissue of the female gametophyte continues to surround the embryo and
serves as nutritive tissue for the embryo upon seed germination (except in angiosperms)
→ The megasporangium (nucellus) eventually degenerates → integument matures into
a peripheral seed coat, which may differentiate into various hard and/or fleshy layers
Note single ring of vascular bundles, with xylem inside, phloem outside. B.
Helianthus stem cross-section, an example of a eustele. C. Close-up of
vascular bundle, showing xylem, phloem, and associated fibers.
SEED ADAPTATIONS
 adaptive significance of the seed
o provide protection, mostly by means of the seed
coat, from mechanical damage, desiccation, and
often predation
o function as the dispersal unit of sexual reproduction
 a fleshy outer seed coat layer may function to
aid in animal dispersal
 differentiation of the seed coat into one or more
wings functions in seed dispersal by wind
o function in dormancy mechanisms that ensure
germination of the seed only under ideal conditions
of temperature, sunlight, or moisture
o upon germination, the nutritive tissue surrounding
the embryo provides energy for the young seedling,
aiding in successful establishment
 (in seed plants) female gametophyte (which develops
within the megaspore) remains attached to and
nutritionally dependent upon the sporophyte
o reverse condition as is found in the liverworts,
hornworts, and mosses
EUSTELE
 eustele – a primary stem vasculature that consists of a
single ring of discrete vascular bundles
o Each vascular bundle contains an internal strand of
xylem and an external strand of phloem that are
radially oriented
 Protoxylem endarch – toward the center of the stem
o Lycophytes: exarch protoxylem
o most monilophytes: mesarch protoxylem
DIVERSITY OF WOODY AND SEED PLANTS
Cycadophyta—Cycads
 Cycadophyta (Cycadales) – MONOphyletic lineage
ARCHEOPTERIS
 Archeopteris – fossil plant and a well-known lignophyte
that lacked seeds
o a large tree, with wood like a conifer but leaves like a
fern
o Sporangia – producing spores, were born on fertile
branch systems
o Some species of Archeopteris were heterosporous
“PTERIDOSPERMS”— “SEED FERNS”
 Pteridosperms (Seed Ferns) - a PARAphyletic group of
fossil plants that had fernlike foliage, yet bore seeds
 Medullosa – a well-known example of a seed fern
o fernlike leaves – in the form genera Alethopteris and
Neuropteris
o pollen-bearing organs – in Dolerotheca, which had
huge pollen grains
o seeds of Medullosa – placed in the form genus
Pachytesta
GYMNOSPERMAE—GYMNOSPERMS
 2 sister groups of Spermatophyta (seed plants)
o Gymnospermae
o Angiospermae
 Gymnospermae (after gymnos, naked + sperm, seed) –
ovules are not enclosed by a surrounding carpel layer
(thus, being “naked”) at the time of pollination
o developing seeds – often enclosed by
megasporophylls or ovuliferous scales, after
pollination
o nonflowering seed plants
 Diversity of Gymnospermae
o cycads (Cycadophyta) – most basal lineage
o Ginkgo (Ginkgophyta)
o conifers (Coniferae)
 Gnetales – either sister to the conifers or within
the conifers, as the sister group to the Pinaceae
consisting of plants with a mostly short, erect stem or
trunk, rarely tall and palmlike (as in Microcycas)
o approximately 320–340 species in 11 genera
o found in southeastern North America, Mexico,
Central America, some Caribbean islands, South
America, eastern and southeastern Asia, Australia,
and parts of Africa
o leaves – spirally arranged, mostly pinnately
compound
 Bowenia – only genus with bipinnately
compound leaves
o loss of axillary branching on the aerial trunk
 no lateral (axillary) branching
o circinate vernation
 cycad pinnae (Cycas) or leaves (some Zamiaceae;
e.g., Bowenia)
o dioecious – either male or female
o pollen cones/strobili (male cones/strobili) – consist
of an axis bearing microsporophylls, each of which
bears numerous microsporangia
 microsporangia – produce many microspores
(n), each of which develops into a pollen grain
 motile sperm cells
o 2 families - differing primarily in the absence and
presence of seed cones
 Cycadaceae – no seed cones
 seeds are produced on the margins of
numerous megasporophylls
 Zamiaceae – have seed [ovulate] cones/strobili
(female cones/strobili)
 Seed cones – consist of an axis bearing
megasporophylls, which bears two seeds
o economic importance
 source of food starch (sometimes termed
“sago”)
 Cycas revoluta (“sago palm”) – planted
horticulturally
 Cycadaceae Zamiaceae
Cycads Family Coontie Family
1 genus (Cycas, incl. 10 genera/ca. 220–230 species
Epicycas)/100–110 species (incl. Boweniaceae, Stangeriaceae)
dioecious trees or perennial herbs dioecious trees or perennial herbs
trunks or subterranean stems trunks or subterranean stems
large, coriaceous, evergreen,
pinnate leaves leaves pinnate [rarely bipinnate]
(vernation involute circinate)
large, determinate pollen cones determinate pollen and seed cones
No seed cones With seed cone
numerous toothed to divided
peltate megasporophylls each
megasporophylls arising from apex
bearing 2 [3] adaxially marginal
of trunk, each bearing one or more
ovules/seed
marginal ovules/seeds
Ginkgophyta
 Ginkgophyta (ginkgophytes) – extensive fossil record but
contain only one extant species, Ginkgo biloba
o Ginkgo biloba – native only to certain remote regions
of China
 a highly branched, woody tree
 short shoots in addition to long shoots
 distinctive obtriangular (fan-shaped), often two-
lobed leaves with dichotomous venation
o dioecious
 Male Ginkgo trees bear “cones” without
sporophyll
 pollen cones – consist of a central axis with
lateral stalks each of which bears two
microsporangia
 ancestrally motile sperm
 Female Ginkgo trees do not bear cones
 female reproductive structures each consist
of an axis having two terminal ovules
 Ginkgoaceae (Maidenhair Tree family)
o resinous, dioecious trees
o roots: mycorrhizal
o stem: trunk is tall, with numerous lateral branches
o leaves: simple, spiral, petiolate, deciduous, and
flabelliform (fan-shaped), with open dichotomous
venation
 born on both long shoots and stout short shoots
o male reproductive structure: catkin-like, each
consisting of an axis bearing microsporophylls with
paired, pendant microsporangia
o ovulate reproductive structure: consisting of an axis
bearing generally two erect ovules each with a basal
collar
o seed – with an outer fleshy and inner hard
integument layer
Coniferae—Conifers
 Coniferae (Pinophyta or Coniferophyta) – ancient group of
land plants that were once dominant in most plant
communities worldwide
o MONOphyletic group of highly branched trees or
shrubs with simple leaves
o Leaves: linear, acicular (needle-like), or subulate (awl-
shaped)
o short shoots – adjacent internodes are very short in
length
 fascicle (in Pinus) – a specialized short shoot
consisting of stem tissue, one or more needle-
shaped leaves, and persistent basal bud scales
o loss of sperm cell motility – apomorphy with
Gnetales
 siphonogamy
 male gametophyte of conifers delivers the sperm
cells more directly to the egg by the growth of
the pollen tube to the archegonial neck
 more than one archegonium per seed, multiple
fertilization events may occur, resulting in
multiple young embryos, but usually only one
survives in the mature seed
o dioecious
o Seed cones – compound structure, consisting of an
axis that bears modified leaves called bract
 ovuliferous scale – separate seed-bearing
structure
 modified lateral shoot system
 Cordaitales (fossil group) – had seed cones that
consisted of an axis bearing bracts
 in the axils of the bracts was a shoot
(branch) system, which consisted of an axis
bearing sterile leaves and one to many
ovules
 compound cone hypothesis – veins of the
ovuliferous scales of extant conifers are inverted
(upside down), with respect to the bract,
indicating that these veins may have been
derived from ancestral leaves oriented 180˚
relative to the bract
 Pseudotsuga (Douglas-fir) – bracts are elongated
and can be seen on the outside of the ovuli
ferous scales
 Podocarpaceae and Taxaceae – seed cones are
greatly reduced
 bracts and ovuliferous scales – only present
as developmental remnants or modified to
form a fleshy structure, with the entire cone
bearing only one ovule
Evolution of the compound conifer cone. A,B. CORDAITALES, Lebachia. A.
Compound cone, bearing numerous, spirally arranged bracts, each subtending
a fertile short shoot. B. Close-up of bract and axillary fertile shoot system (left
= abaxial view, right = side view), bearing spiral leaves and single ovule. C,D.
Fossil conifers. C. Voltzia, shoot system of compound cone, with fi ve sterile
leaves and three ovules. D. Ulmannia, shoot system of compound cone (left =
abaxial view, right = adaxial view), showing outer bract and inner fused shoot
system (ovuliferous scale) and ovule.
 Chapter 6
Evolution of Flowering Plants
 Angiosperms (Flowering plants, Magnoliophyta, or
Anthophyta) – MONOphyletic group currently thought to
be the sister group to the gymnosperms
o 95% of all land plant species – most numerous,
diverse, and “successful” extant plant group
ANGIOSPERM APOMORPHIES
(1) FLOWER
 Flower – a modified, determinate shoot system bearing
one or more stamens, collectively called the androecium,
and/or one or more carpels (making up one or more
pistils), collectively called the gynoecium
o bisexual (perfect) – both stamens and carpels
o unisexual (imperfect) – only stamens or carpels
 perianth (tepal) – consisting of modified leaves at the base
of the shoot system
o protects the other floral parts during floral
development and an attractant for pollination
o perianth of two discrete whorls or series of parts:
 an outer calyx (outer tepal) and an inner corolla
(inner tepal)
 calyx – green and photosynthetic, composed of
leaf-like sepals or calyx lobes (if fused)
 corolla – typically colorful, showy, and
odoriferous and is composed of individual petals
or corolla lobes (if fused)
o tepals/perianth parts/perianth segments – spirally
arranged units with no clear point of differentiation
 primordia – bumplike mounds of tissue formed by actively
dividing regions of cells
o develop in whorls from outside to inside
 sepal (outer tepal) primordia → petal (inner
tepal) primordia → stamen primordia (often in
two or more whorls) → carpel primordia
o floral development – homologous to leaves
 initiating like leaf primordia of a vegetative shoot
 being innervated by veins
 often having a dorsiventral shape
 “ABC” model of floral development
o Arabidopsis thaliana (Drosophila of the plant world)
o Sepals – by A
o Petals – by A and B
o Stamens – by B and C
o Carpels – by C
o SEPALLATA class – needed in combination with those
of the A, B, and C classes to effect proper floral organ
identity
 transcription factors – induce the expression of other
genes that bring about the development of the 4 floral
organs
 Animal pollination – appears to be the primitive condition
in the angiosperms, separating them from the
predominantly wind-pollinated gymnosperms
o may include bees, butterflies and moths, flies, bats,
and birds
 Ranunculus sp.(buttercup) – bee-pollinated
 Calonyction sp. (moon flower) – moth-pollinated
 Stapelia sp. (star flower) – fly-pollinated
 Selenicereus (night-blooming cereus) &
Couroupita guianensis (cannonball tree) – bat-
pollinated
 Strelitzia reginae (bird of paradise) – bird-
pollinated
 water pollinated or wind pollinated – flowers of many
groups that are quite reduced in size or structural
complexity, often lacking a perianth altogether
o Phyllospadix torreyi (surf-grass) – water-pollinated
o Grass – wind-pollinated
(2) STAMENS
 stamen – male reproductive organ of a flower
o modified microsporophylls, modified leaves that bear
microsporangia
o Microsporangia → microspores → pollen grains
 laminar (leaf-like) structure (in some stamen) – to which
the anther is attached or embedded
o sessile – Some stamens that lack a filament (or
lamina) and is directly attached to the rest of the
flower
 stamens of most flowering plants have two parts
o filament – stalk
o anther – pollen bearing part
 Anther
o synangium – a fusion product of sporangia
o unique in (ancestrally) containing two pairs of
microsporangia, usually arranged in a bilateral
symmetry
o theca – discrete half of the anther which contains
each pair of microsporangia
 bithecal – two thecae
 tetrasporangiate – 4 microsporangia
o anther locule – single, contiguous chamber from the
2 microsporangia that coalesced at maturity
o ** anthers of some angiosperms are monothecal or
bisporangiate (secondarily reduced to a single theca)
 adaptive value of the stamens of angiosperms over the
microsporophylls of gymnosperms – selective pressures
for the flower
o smaller & lighter than gymnosperm microsporophylls
o occur in bisexual flowers, rather than in more
massive, unisexual cones
o enabled the evolution of specialized pollination
mechanisms
(3) REDUCED MALE GAMETOPHYTE
 reduced, 3-celled male gametophyte
o microspore - formed by meiosis within the
microsporangium
 single nucleus divides mitotically to form 2 cells:
tube cell & generative cell
 microspore → pollen grain
o pollen grain – immature, endosporic male
gametophyte
 generative cell divides one time – producing two
sperm cells
 Pollen Tube
o siphonogamy – elongate pollen tube grows out of the
pollen grain wall due to germination of pollen grains
o (In gymnosperms) the pollen tube develops after the
pollen grains enter the micropyle of the ovule and
functions as a haustorial device for a long period of
time
o (In angiosperms) the pollen tube forms immediately
after transfer of pollen to the stigma
 pollen tube (angiosperms) elongates through (and feeds
upon) the tissues of the stigma and style of the carpel →
reaches the ovule → penetrates the micropyle →
transports the two sperm cells directly to the female
gametophyte
o NON-motile sperm – conifers and angiosperms
 adaptive significance of the reduced male gametophytes
of angiosperm – evolution of a reduced female
gametophyte and relatively rapid seed development
o to deliver sperm cells to the female gametophyte and
effect fertilization very rapidly than in gymnosperms
(4) CARPEL
 carpel – major apomorphy of angiosperms
o constitutes a modified, conduplicate megasporophyll
bearing two, adaxial rows of ovules
 megasporophyll – a modified leaf that bears
megasporangia, which in the seed plants are
components of the ovules and seeds
 Conduplicate – inwardly folded longitudinally
and along the central margin
o At maturity the carpel body completely encloses the
ovules and seeds, accounting for the name
angiosperm (Gr. angio, vessel + sperm, seed)
 sporophyll-like nature of the carpel
o develop like a leaf – initially flattened, dorsiventral
shape, with an adaxial (toward the top-center) and
abaxial (away from the top-center) surface
o has veins
 dorsal (median) vein or bundle – one in the
middle, corresponding to the midvein of a leaf
 ventral (lateral or placental) veins/bundles –
two others near the two carpel margins
 ascidiate carpel – develops from a ring of tissue that
grows upward, sometimes assuming a peltate form
 flowers with 2 or more carpels:
o apocarpous (distinct) – separate from one another
o syncarpous (connate) – fused together
 gynoecium – totality of female reproductive structures in a
flower, regardless of their structure
o carpel may be alternatively defined as a unit of the
gynoecium
o composed of one or more pistils with basal ovary, an
apical style (or styles), which may be absent, and one
or more stigmas, the tissue receptive to pollen grains
 simple pistil – pistil equivalent to one carpel
 compound pistil – composed of 2 or more fused
carpel
 adaptive significance of carpels
o carpels – receivers of pollen
 function to selectively control fertilization
 incompatibility reaction – chemicals that
are present in the stigma and style may
inhibit either pollen germination or pollen
tube growth
 mediated by incompatibility genes
 may inhibit inbreeding, allowing for
reproduction only between genetically
dissimilar individuals of the species
o fruit formation and seed dispersal
 fruit – mature ovary or ovaries (made up of one
or more carpels) plus any accessory tissue that
might be present
 pericarp – mature ovary wall
 seed dispersal
 if the pericarp is fleshy, fruits are dispersed
by animals
 by wind – aided by the development of
wings or trichomes
 by water – via various flotation devices
 mechanically – by various explosive,
hygroscopic, or catapulting methods
 monosporic, 4-nucleate and celled condition – ancestral
condition found in virtually all Nymphaeales and
Austrobaileyales
o Nuphar/Schisandra type
o ancestral module – one polar nucleus in a central cell
and 3 cells (the egg apparatus) at the micropylar end
 doubled (a third sequence of mitotic divisions) to
yield the common Polygonum type
 quadrupled to yield something like the 16-
nucleate Penaea type
 Amborella trichopoda – most basal angiosperm
o Amborella type – modified type of female
gametophyte, being 9-nucleate and 8-celled via an
extra mitotic division in the egg apparatus producing
a third synergid cell

(5) TWO INTEGUMENTS

 bitegmic ovules – growth of two integuments during ovule

development
o unique apomorphy of angiosperms
o unitegmic – all nonflowering seed plants have ovules
with a single integument
o two integuments typically coalesce during seed coat
development, but may form anatomically different
layers

 (unclear) adaptive significance of two integuments

o enabled the evolution of specialized seed coat layers

(6) REDUCED FEMALE GAMETOPHYTE

 reduced female gametophyte – major apomorphy

o (seed plants) a single megasporocyte within the
megasporangium (nucellus) divides meiotically to
form four haploid megaspores
 female gametophyte typically generates from
only one of these megaspores
o (angiosperms) megaspore divides in a sequence of
three mitotic divisions, resulting in a total of eight
haploid nuclei
 adaptive significance of a reduced female gametophyte -
developmental timing
o angiosperms have the capacity to more quickly
generate seeds
o evolution of rapidly spreading annual herbs
(7) ENDOSPERM FORMATION

 Polygonum type – pattern of arrangement of eight nuclei  endosperm – major apomorphy

into seven cells
o in micropylar region, 3 cells develop: an egg cell
flanked by two synergid cells
 egg apparatus = egg cell + synergid cells
o In the chalazal region, 3 antipodal cells form
o central cell – remaining volume of female
gamethophyte which contains two polar nuclei
 “embryo sac” – female gametophyte of angiosperms
o ** Archegonia do not form within the female
gametophyte of angiosperms
o product of double fertilization
o pollen tube → micropyle of the ovule, penetrates one
of the synergid cells and releases the two sperm cells
into the central cell of the female gametophyte
 One sperm cell migrates toward and fuses with
the egg cell to produce a diploid zygote
 other sperm cell fuses with the two polar nuclei
to produce a triploid, or 3n, endosperm cell
 endosperm – mass of tissue that generally
envelopes the embryo of the seed from the
repeated mitotic division of endosperm cell
 endosperm – replaces the female gametophyte as the
primary nutritive tissue for the embryo in all angiosperms
o containing cells rich in carbohydrates, oil, or protein
 adaptive significance of endosperm – developmental
timing
o endospermous nutritive tissue of angiosperms does
not begin to develop until after fertilization is
achieved
 in gymnosperms, considerable female
gametophytic nutritive tissue is deposited after
pollination, even if the ovules are never
ultimately fertilized
o conservation of resources, such that seed storage
compounds are not formed unless fertilization is
assured
o triploid - Having three sets of chromosomes (one
from the male and two from the female) may enable
ORIGIN OF ANGIOSPERMS
the endosperm to develop more rapidly (correlated
with rapid overall seed development) and may also
provide greater potential for chemical variation in
nutritive contents
(8) SIEVE TUBE MEMBERS
 sieve tube members – specialized sugar-conducting cells
o Sieve cells (and associated albuminous cells) –
primitive sugar-conducting cells and are found in all
nonflowering vascular plants
 evolutionarily modified from sieve cells and are
found only in flowering plants
 difference: pores at the end walls are
differentiated, being much larger than those on
the side walls
 sieve plates – collections of differentiated
pores at the end walls
 compound (composed of two or more
aggregations of pores)
 simple (composed of one pore region)
o companion cells – Parenchyma cells associated with
sieve tube members
 function to load and unload sugars into the
cavity of sieve tube members
 adaptive signifi cance of sieve tube members over sieve
cells - may provide more efficient sugar conduction
VESSELS
 One angiosperm specialization concerns water and
mineral conductive cells
 vessels – two ends of the cells have openings, termed
perforation plates
o Amborellales, some Nymphaeales – vesselless, having
only tracheids (which lack perforation plates)
 Specializations of vessels
o modification of the perforation plate from scalariform
to one with fewer, less transversely oriented
openings, to a simple perforation plate (having a
single opening
o modification from tapering end walls to
perpendicular ones
o modifIcation from long, narrow cells to short, wide
cells
 adaptive advantage of vessels over tracheids
o more efficient solute conduction due to greater rate
and lower flow resistance
o under conditions of low water potential or freezing
temperatures, wider vessels may increase the chance
of cavitation (formation of a gas bubble in the water
stream), which interrupts water fl ow and can result
in plant death if not self-repaired. This may explain
why tracheid-bearing conifers dominate over
angiosperms in cold climatic regions
 Charles Darwin: “abominable mystery”- relatively rapid
diversification of the “higher plants” (presumed to mean
angiosperms)
 fossil records
o dispersed pollen grains from Cretaceous period, 140
million years ago – earliest definitive fossil
o earliest definitive flowers – as early as 130 million
years ago
 Amborella trichopoda of the Amborellales – most basal
angiosperm lineage
o lacks vessels
o unisexual flowers with a spiral perianth, laminar
stamens, and separate carpels
 “pteridosperms” – a paraphyletic assemblage of extinct
plants that possessed seeds and had generally fernlike
foliage
o possible angiosperm progenitors
o Caytonia of the Caytoniales – fossil taxon that
exemplifi es a putative transition to angiosperms
 possessed reproductive structures similar to
those of the angiosperms
 male reproductive structures resemble
anthers in consisting of a fusion product
(synangium) of 3 or 4 microsporangia;
 radially (not bilaterally) symmetric
 female reproductive structures consist of a
spikelike arrangement of units, cupules
 cupule = carpel
 ancestral carpel morphology, a
conduplicate megasporophyll bearing
ovules along two margins
 cupule did not function as a carpel in
terms of a site for pollen germination
o Glossopteridales or Glossopterids – trees with simple
leaves having a midrib giving rise to an extensive
reticulate venation system
 open, leaf-like capitulum may represent a
partially closed megasporophyll, transitional to
an angiospermous carpel in which the
megasporophyll encloses ovules
 genus Archaefructus – have bona fide carpels,