Vascular Plant Apormophies

Vascular = Tracheophytes

 Monophyletic subgroup of the land plants.

 Vascular plants major apomorphies
→ Independent, long-lived sporophyte
→ Branched sporophyte
→ Lignified secondary walls
→ Sclerenchyma
→ Tracheary elements – xylem
→ Sieve elements – phloem
→ Endodermis
→ Roots
Unlike the Bryophytes…
→ Vascular plants, have a dominant, freeliving, photosynthetic, relatively persistent sporophyte
generation
→ Gametophyte – is free living; haploid
 Photosynthetic
 Smaller and short lived than the sporophyte
Branched Sporophyte
→ Stems are different from the Bryophytes
→ Branched and bear multiple sporangia
→ Polysporangiomorpha
 This apomorphy is shared among transitional between the “bryophytes” and the
tracheophytes
→ Earliest vascular plant stem – dichotomous
 Splits into two
 Have equal meristems; grows independently
→ Later lineages evolved the pseudomonopodial
 Starts out dichotomous
 Then one branch becomes dominant
→ Sporophytic stems of vascular plants function as:
 Supportive organs
 Bearing and elevating reproductive organs and leaves
 Conductive organs – via vascular tissue
Lignified Secondary Cell Walls
→ Vascular plants posses a chemical known as lignin,
 Which is added to the secondary cell wall; imparts additional strength
 Found in certain specialized cells of vascular plants
→ Secondary cell walls are secreted to the outside of the plasma membrane; between primary
cell wall
 Secreted when cells have stopped elongating
LYCOPODIOPHYTA – Lycophytes
→ Roots
a. Dichopodial – the root apical
meristem may branch into two roots.
b. No lateral roots develop
c. endarch protoxylem – protoxylem
present towards the center or interior of
the metaxylem.
→ Stem have an exarch protoxylem
→ Ancestral sporangia
a. dorsiventral – flattened and having a
dorsal, upper, and vetral, lower,
surface. Distinct or dissimilar upper and
lower faces or surfaces.
b. dehisces transversely relative to the
axis of the stem or subtending leaf.
→ Leaves
a. Lycophylls – sporophytic leaves
called “leaves”.
a.1. Single unbranched vein
a.2. arise from the intercalary
meristem
a.3. Lacking leaf gap – central
vascular tissues are continuous and
uninterrupted
3 Extant families:
1. Lycopodiaceae – Lycopodiopsida
(club-mosses)
2. Isoetaceae – Isoetes
(Quillwort family)
3. Selaginellaceae – Selaginella
(Spike-Moss family)
EUPHYLLOPHYTA – Euphyllophytes
→ Roots
a. Monopodial – they do not dichotomously branch at
the apical meristem.
b. Have lateral roots – arise endogenously from either
the endodermis and pericycle
c. exarch protoxylem – protoxylem is positioned
towards the periphery or exterior of the metaxylem
where in the metaxylem towards the center.
→ Ancestral sporangia
a. terminal in position
b. longitudinal dehiscence
→ 30-kilobase inversion
→ Leaves
a. Euphylls
a.1. multiple veins or are highly branched
system of veins
a.2. arise from the marginal or apical
meristem
a.3. Associated with leaf gap – parenchyma
tissue interrupting the vasculature of the
stem.
2 major groups, which are sister to one another:
1. Monilophyta – Monilophytes (Ferns)
2. Spermatophytes (Seed plants)
LYCOPODIOPHYTA – LYCOPHYTES
Lycopodiopsida (Club-mosses)
Apomorphies
1. Homospory – having one type of spore. Production of morphologically identical spores (asexual)
2. Some members may resemble a moss
3. Sporangia develop laterally in the axils of sporophylls
is a leaf that produces or bears the spores.
The sporohpylls in some family members are:
A.) Similar to vegetative leaves Co-occur with them on shoots that are
indeterminate continuous growth
OR
B.) Vary in size and are aggregated into a terminal shoot system that are
determinate with associated sporangia, known as strobilus or cone
Terminates growth after formation. a modified,
determinate,
reproductive shoot
system, consisting of a
stem axis bearing
sporophylls.
Lycopodiaceae – (Club-mosses family)
→ are distinctive in being homosporous, dichotomously branched, erect, prostrate, or
pendulous, perennial, lycophyllous herbs
→ Roots – adventitious, endogenous, dichopodial
grows from the pericycle
are plant roots that form from any nonroot tissue; arising from the
stem of the plants
→ Stems – dichotomously branched rhizomes
→ Leaves – simple, sessile, spiral or whorled
→ Sporangia – homosporous; occurring on short stalks in axils of leaves
→ Gametophytes are mycorrhizal
Fungal roots
Mycorrhizae are fungi that have symbiotic relationship with the
roots. Their major role is to enhance nutrient or water uptake of the
host (roots) by expanding a large volume of soil that roots alone
can do. The fungus receives its necessary carbohydrates from the
host plant as it cannot photosynthesize, therefore cannot fix its
carbon. In return, the mycorrhizae absorb nutrient from the soil
which are passed along the plant.
Isoetopsida
 Isoetaceae
 Selaginellaceae
Isoetopsida differ from those of the Lycopodiopsida in having:
→ Leaf ligules – appendages on the upper side of the leaf; membranous appendage
growing from the inner side of the leaf near the junction with the leaf sheath.
→ Hydrates young, developing lycophylls.
→ Heterosporous – refers to the production of two types of spores:
→ Microspores – small and numerous
→ Megaspores – large and fewer
* In both Selaginella and Isoetes:
→ megaspore develops into a female gametophyte- archegonia
→ Each microspore germinates to form a male gametophyte- antheridia
→ Spores are endosporic – gametophytes develop entirely within the original spore wall

Isoetaceae – Quillwort Family aquatic or terrestrial perennial herbs

can live for several years
→ The Isoetaceae are distinctive in being cormose to meizomatous plants
→ with a basal rosette of microphyllous, ligulate leaves
one single unbranched vein
→ The leaves basally sheathing, apically linear to acicular
→ Heterosporous leaves, bearing adaxial megasporangia or microsporangia within
sheathing leaf base.
→ Gametophytes are endosporic

Selaginellaceae – Spike-moss family

→ are distinctive in being erect to prostrate herbs
→ with dichotomously branched stems; sometimes form a planar branch system
→ the leaves microphyllous, spiral, either homomorphic or dimorphic and four-rowed
→ sporangia heterosporous; microsporangia and megasporangia born in axils of ligulate
sporophylls of terminal strobili;
→ gametophytes endosporic.

Alternation of generation
There is a multicellular haploid stage as well as a multicellular diploid stage. The microspore
will develop into a male gametophyte (n) antheridium and the megaspore will develop into a
female gametophyte (n) archegonium that produces the gametes, sperms and eggs, respectively,
through mitosis. Then, through fertilization, they form a diploid zygote. The zygote will go through
numerous rounds of mitosis and develop a sporophyte (2n), a fully developed sporophyte body that
will produce the spores. The spores will further differentiate and become more specialized into
microsporocytes and megasporocytes. Through meiosis, they will germinate into a new
gametophyte (n), and thus the alternating cycle is complete.
EUPHYLLOPHYTA - EUPHYLLOPHYTES
Monilophyta – Monilophytes (Ferns)
Four major lineages:
1. Equisetopsida
(Horsetails)
2. Psilotopsida
(Whisk ferns and Ophioglossoids)
3. Marattiopsida
(Marattioid ferns)
4. Leptosporangiatae
(Leptosporangiate ferns)

Apomorphies
→ Siphonostele – a stem vasculature; a ring of xylem is surrounded by an outer layer of phloem.
a. Ectophloic – outer layer of phloem. A type of siphonostele where the xylem is
surrounded by the phloem cylinder on the outer side.
b. Amphiphloic – surrounded on both outer and inner layer of phloem. Phloem is both
around the external of the xylem and the internal of the xylem ring.
c. Dictyostele – a dissected siphonostele. Several leaf gaps separate a single ring of
xylem. This xylem ring is surrounded by phloem.
→ Stem protoxylem is mesarch – protoxylem is in middle of the metaxylem. Tracheary elements first
mature near middle of a patch of xylem; this protoxylem is restricted to the lobes of the xylem.

Equisetopsida – Equisetophytes, sphenophytes, sphenopsids

(Horsetails)

Apomorphies
1. Ridged stems – often associated with hollow canals
2. Reduced, whorled leaves that are usually marginally fused
3. Sporangiophores, consists of peltate axis bearing pendant longitudinally dehiscent
sporangia.
A sporangia-bearing receptacle found in ferns. A scale in horsetails.
4. Photosynthetic spores with elaters
to increase dispersal because they push the spores out of
the plant by absorbing moisture.

Family Equisetaceae
→ Readily distinguished in being rhizomatous, perennial herbs,
→ aerial shoots hollow, ridged, with siliceous epidermal cells and internal canals,
→ leaves small, nonphotosynthetic, simple, microphyllous, whorled, and laterally connate
(sheathed), the apices teeth-like;
→ sporangia in terminal strobili, homosporous, several born underneath peltate
sporangiophores
→ spores green, with four, spatulate, hygroscopic elaters.
PSILOTOPSIDA
Ophioglossales – Ophioglossales Ferns or Ophioglossoid Ferns
→ Leaves have a sterile segment- the photosynthetic blade
→ leaves that are capable of photosynthesis
→ fertile segment- bearing the sporangia
2 jobs of the leaves of the ophioglossales seperately:
1. There is the sterile segment that does the photosynthesis.
2. There is the fertile segment that does the production of spores.

→ Eusporangia- arise from several cells

Ophioglossaceae – Adder’s Tongue family

→ homosporous, eusporangiate, perennial herbs,
having sporangia developing from several epidermal cells rather
than a single cell.
→ leaves simple to compound, bearing a stalked fertile segment containing eusporangia,
→ The gametophytes nonphotosynthetic.

Psilotales – Psylotophytes or Whisk Ferns

Consist of only two genera plants:
1. Psilotum
2. Tmesipteris
→ Lack true roots – instead have rhizoids believed to a primitive retention.
functions like a root in support or absorption.
A short, thin filament or little hair-like
structures.
→ Enations- unlike leaves, they lack vascular tissue; an outgrowth.
→ Synangium- fusion product of two or three sporangia.
Psilotaceae
→ distinctive in being rootless, rhizomatous, perennial herbs, with unbranched or
dichotomously branched stems,
→ aerial stems photosynthetic,
→ the leaves simple, spiral or distichous, lacking vasculature or with a single vein
(microphyllous)
a leaf with single unbranched veins and no discernible gap around the leaf
trace.
→ sporangia 2- or 3-synangiate, born on a short lateral axis subtended by a bifid
appendage,
arising from the stem
→ the gametophytes.
Marattiopsida – Marattioid Ferns
Contains one single order Marattiales and Family Marattiaceae.
Very similar to Polypodiopsida and Leptosporangiate ferns.
Apomorphies
→ Eusporangiate- whose sporangia arise from several epidermal cells
→ Synangium- fusion product of two or three sporangia
→ A distinctive apomorphy of the Marattiales is the occurrence of a “polycyclic
siphonostele”.

*Eusporangiate Ferns – vascular spore plants produce sporangia from several epidermal
cells rather than a single cell.

Marattiaceae – Marattia Family

→ distinctive in being large, terrestrial ferns with mucilage canals,
role in the storage of
water and food, seed
germination, and
thickening membranes.
→ the stems with a polycyclic dictyostele,
numerous xylem rings, each with several leaf
gaps. Each ring's centripetal side contains
phloem.
→ the leaves generally large, simple to several-pinnate,
→ with abaxial, intramarginal eusporangia, sometimes fused into synangia.
Bottom part of the leaf
Polypodiopsida – Leptosporangiate Ferns
Of the major monilophyte groups, the leptosporangiate ferns contain by far the greatest diversity,
estimated from at least 8800 to over 12,000 species.

Apomorphy
 Leptosporangia are unique among vascular plants in
(1) developing from a single cell
(2) having a single layer of cells making up the sporangium wall.

 much smaller spore number than eusporangia

*receptacle – where the leptosporangia are attched
*Indusium- a thin membranous covering, especially a shield covering a sorus on a fern frond.

A) Rhizome- horizontal underground plant stem

B) Pinnae- first leaf discrete leaflets

 The ultimate leaflets or blade divisions are called pinnules.

D) Fern petiole is a stalk that attaches a leaf to the plant stem

 covered with scales

E) Croziers or fiddleheads – undergoes Circinate vernation in which coiled early in development and
uncoil at maturity. the development of a frond from a fern fiddlehead.
 Evolution and Diversity of Woody and Seed plants
LIGNOPHYTA – Lygnophytes (Woody plants)

 Monophyletic lineage of euphyllous vascular plants

 Vascular cambium
 Cork cambium
 Heterospory
 Seed
 Pollination droplet
 Endosporic male gametophyte
 Eustele

 Vascular cambium – cells that develops within the stems and roots as a continuous layer, between the
xylem and phloem
→ Gives rise to wood, and a cork cambium
produces cork
→ Arise during secondary growth

Functions:
 Precursor to the formation of intricately branched shrubs or tree with tall overstory canopies
 Continues to divide into:
→ secondary xylem (wood)- produced to the inside of it
Functions:
 Enables plant to grow tall
 To acquire systems of lateral branches
 Structural support
→ secondary phloem-produced to the outside
Functions:
 Protects from mechanical damage, predation, and desiccation
extreme dryness
→ Bifacials- layers are produced in both sides
→ Generally, much more secondary xylem is produced than the secondary phloem
 Cork cambium differentiates at the periphery
→ Cork cambium and derivatives- periderm
→ The outermost layer is the cork
Cork functions:
 Contains a polymer of suberin to prevent water loss
 Thick cells that protects the delicate vascular cambium
 lignophytes also undergo monopodial growth
→ Single main shoot develops
→ lateral branches grow from axillary buds
 SPERMATOPHYTES – SEED PLANTS
 A monophyletic lineage within the lignophytes
o The major novelty that unites this group is the seed
 Seed- An embryo, an immature diploid sporophyte developing from the zygote
o surrounded by nutritive tissue and enveloped by a seed coat
 Radicle – immature root
 Epicotyl – Shoot Apical Meristem
 Cotyledons – young seed leaves
 Hypocotyl – transition between root and stem

SEED EVOLUTION
1) Heterospory-formation of two types of haploid spores within two types of sporangia:
 megasporangium ->megaspores- > Female gametophyte- archegonia
 microsporangium- >microspores- > Male gametophyte- antheridia
 heterospory has evolved independently in other, nonseed plants
o e.g lycophytes

2) Endospory- complete development of the female gametophyte within the spore wall
 Exospory was the ancestral trait
external growth of female gametophyte
3) Reduction of megaspores number to one
 four haploid megaspores produced by meiosis
→ three consistently abort, leaving only one functional megaspore.
→ More space; can increases in size
4) Retention of the megaspore
 Instead of being released from the megasporangium (ancestral trait)
o This was accompanied by a reduction in thickness of the megaspore
wall
5) Evolution of the integument & micropyle
 final event in seed evolution
 An Integument envelope the megasporangium
→ Likely evolved from the telomes
 Lagenostome – functions to funnel pollen grains to a pollination chamber
 Micropyle – replaced the ancestral lagenostome
Facilitates the entry of pollen into ovary
Functions in controlling the water absorption during the seed germination

POLLINATION DROPLET
 Evolutionary novelty associated with seed evolution
 A droplet of liquid that is secreted by young ovule through the micropyle.
→ Contains water plus some sugars or amino acids
o formed by the breakdown of cells at the distal end of the
megasporangium
 The pollination droplet transports pollen grains through the micropyle.
→ the resorption of droplet pulls the pollen grain into the pollination chamber
POLLEN GRAIN
• Concomitant with the evolution of seeds
• pollen grains = “immature” endosporic male gametophyte
o extremely reduced male gametophytes
• Development of the male gametophyte within the original spore wall.
• Needs to be transported to the micropyle of the ovule
• Male gametophyte grows an exosporic pollen tube
• Functions as a haustorial organ
Penetrate the surrounding sporohytic tissue
and obtains nutrients by absoption.

POLLEN TUBE
 Male gametophytes of all extant seed plants form a pollen tube soon after the pollen grains make
contact with the microsporangial tissue of the ovule.
 Motile sperm is delivered into a fertilization chamber, goes to the archegonium containing the
egg, a process known as:
• zooidogamy
 Conifers do siphonogamy- deliver nonmotile sperm cells to the archegonium

SEED ADAPTATIONS
 Protection – By means of the seed coat
 Dispersal unit – Animals can disperse them
 Dormancy mechanisms – ensures to germinate under ideal conditions
 Nutritive tissue – Surrounds the embryo and nourishes it

EUSTELE
→ Consists of a single ring of discrete vascular bundles that contains:
o Internal strand of xylem
o External strand of phloem
o Radially positioned
 A eustele is a primary (prior to 2ndary growth) stem vasculature that consists of a single ring of
discrete vascular bundles.

 Eustele is endarch in position