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Evolution of vascular plants

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Plants had an aquatic ancestor probably a Coleochaete-like alga, of the Chlorophyta.
Plant evolution shows a tendency toward greater independence from water as they progressively occupied the land.
Air is drier than water and less buoyant. Land plants had to develop adaptations to conserve and transport water with its solutes, absorb water from the environment, support itself facing the direction of sunlight, solve reproductive problems like fertilization and nourishment of embryo, and dispersal of offspring.

  1. Dominant sporophyte and reduced gametophyte.

  • In bryophytes, the gametophyte is dominant generation.

  • Water is required for fertilization.

  • Pollen and embryo sac are much reduced gametophytes.

  • The occupation of the land by the bryophytes was undertaken with emphasis on the gamete-producing generation, which requires water for fertilization.

  1. Development of fluid-transport system, the xylem and phloem.

  • Aquatic plants take water throughout their entire body.

  • On land, soil is the water reservoir; the air is dry in comparison to cells.

  1. The ability to synthesize lignin.

  • Early land plants were small and probably stayed upright by means of turgor pressure.

  • Lignin adds rigidity to the cell wall and allows the plant to reach greater heights.

  1. Development of apical meristems.

  • Bryophyte sporophyte growth is subapical and unbranched.

  • It allows the sporophyte to branch many times.

  1. Ability to produce many sporangia.

  • Only one sporangium is produced the bryophyte sporophyte.

  • The many branches of vascular plants became capable of bearing many sporangia.

  1. More diverse plant body through the development of roots stems and leaves.

  • Roots for absorption, storage and anchorage.

  • Stems for support above ground, transport and growth toward the light.

  • Leaves for photosynthesis.

  1. Evolution of seeds.

  • Provides the embryo with food and protection.

  • Dispersal of the species to new locations.

The plant body consists of a root system and a shoot system.

  • Root system absorbs and anchors.

  • Shoot system is involved in photosynthesis and its supporting activities.

Cells are organized into tissues and tissue systems.

There are three tissue systems in plants, which occur in all organs of plants and are continuous from organ to organ.
It reveals the basic unity of the plant body.

  1. Dermal tissue provides the protective covering of the body.

  2. Vascular tissue transports materials within the plant body; it is embedded in the ground tissue.

  3. Ground tissue fills in the spaces between the other tissues and stores water and food.

Primary growth is produced by the activity of the apical meristems.
The primary tissues that arise from the apical meristems are part of the plant body, the primary plant body.
Secondary growth results from the activity of lateral meristems.

  • Vascular cambium produces vascular tissue every year.

  • Cork cambium forms the periderm, which is mostly cork tissue.

  • Secondary vascular tissue and periderm constitute the secondary plant body.

The periderm eventually replaces the epidermis in older portions of the plant.

The transporting cells of the vascular tissue are...

  • Tracheids and vessel elements in the xylem, also called tracheary elements.

  • Sieve tube members in the phloem.

Sieve tube members have a thin cell wall and do not preserve well as fossil.

Tracheids have a distinctive lignified wall thickening and preserve well in the fossil record.
The earliest vascular plants of the Silurian and Devonian have tracheids that conducted material and provided support
Vessel elements are more specialized than tracheids and are found only angiosperms and gnetophytes.
Vessels probably evolved from tracheids in different groups of plants and are an example of convergent evolution.
The primary vascular tissue and associated ground tissue exhibit three basic arrangements:

  1. The protostele: a solid core of vascular tissue surrounded by ground tissue.

  1. The siphonostele: a pit surrounded by vascular tissue.

  • It has characteristic leaf gaps associated with leaf traces.

  1. The eustele: a system of strands surrounded the pith and separated from one another by ground tissue.

  1. For a more complete description of other types of steles visit:

Roots are derived from the subterranean portion of the ancient stems.

Roots have retained many of the ancient structural characteristics no longer present in stems.
From an evolutionary perspective, there are two kinds of leaves:

  • Microphylls are single veined leaves associated with leaf gaps and evolved as superficial outgrowth of the stem.

Microphylls are associated with protosteles and are found in the lycophytes.

  • Megaphylls have a complex venation pattern, and evolved from a branch system.

Megaphylls are associated with siphonosteles and eusteles.

In ferns, megaphylls are associated with leaf gaps.
The Telome Theory proposed by Zimmerman explains the development of megaphylls from a branch system.

All vascular plants are oogamous.
They have alternation of heteromorphic generations.
Homosporous plants produce one kind of spores; heterosporous plants produce two kinds of spores.
Homosporous plants produce bisexual gametophytes that bear antheridia and archegonia.
Homospory is found psilophytes, sphenophytes, some lycophytes and most ferns.

Heterospory is the production of two types of spores in two types of sporangia.
Heterospory is found in the Selaginellaceae, Isoetaceae and aquatic ferns.
Heterospory was already common in the Late Devonian, about 370 million years ago.

  • Microsporangia produce microspores and give rise to microgametophytes (male).

  • Megasporangia produce megaspores and give rise to megagametophyte (female).

  • These gametophytes are much reduced in size and develop within the spore wall.

The gametophytes of homosporous plants are relatively large and independent of the sporophyte.

The gametophytes of psilophytes and some lycophytes are subterranean and heterotrophic; they depend on endomycorrhizal fungi for nutrition.
Other species are photosynthetic.
There is an overall trend toward reduction of the gametophyte.
In angiosperms, the megagametophyte is reduced to seven cells and the microgametophyte to three cells, and two of them are sperms.
Archegonia and antheridia have been lost in the lineage leading to angiosperms.
Some gymnosperms have archegonia but lack antheridia.
All of the seedless vascular plants have motile sperms and depend on water for fertilization.
Gymnosperms and angiosperms, the seed bearing plants, depend on pollination prior to fertilization.
Pollination is the transfer of pollen to the vicinity of the megagametophyte.
Pollen grains produce a pollen tube through which motile sperms (cycads and Ginkgo) swim or nonmotile sperm are transferred to the egg.

There are three phyla of extinct seedless vascular plants: Rhyniophyta, Zosterophyllophyta and Trimerophytophyta.
The genera Rhynia, Zosterophyllum and Trimerophyton are members of these phyla.
The earliest known go back about 425 million years ago and most went extinct by the end of the Devonian about 370 million years ago.
These three groups were the dominant vegetation from the mid-Silurian to the mid-Devonian, 425 to 370 million years ago.
For the most part they were relatively simple plants 18 in to 36 inches tall. They had the following characteristics:
1. Naked photosynthetic stems

2. Terminal sporangia (some lateral)

3. No roots or leaves

4. They were all homosporous

5. They had protosteles
Pteridophytes, lycophytes and progymnosperms are more complex groups that were dominant from the Late Devonian through the Carboniferous, from about 370 to 290 million years ago.
Seed plants arose starting in the Late Devonian period, about 380 million years ago, and evolved many new lines by the Permian (290-248 million years ago).
Gymnosperms dominated the land floras throughout the Mesozoic until about 100 million years ago.
Angiosperms appeared in the fossil record about 125 million years ago. It became the dominant group about 30 – 40 million years ago, and has remained so until the present.
Phylum Rhyniophyta
The Rhyniophytes are a group of early land plants originally described from the Rhynie Chert, Scotland.
Rhynia appeared in the mid-Silurian record about 425 million years ago.
It became extinct in the mid-Devonian about 380 million years ago.

  • Seedless; produced spores.

  • Dichotomous branching.

  • Terminal sporangia.

  • Homosporous

  • Plant body was not differentiated into roots, stems and leaves.

  • Underground rhizome with rhizoids.

  • Protostele consisting of a core of xylem surrounded by one or two layers of phloem cells.

  • Some had conducting cells similar to hydroids rather tracheids; they are called protracheophytes.

  • There is evidence of isomorphic alternation of generations.

  • Examples: Rhynia, Cooksonia, Aglaophyton.

Fig. 17-12 is the reconstruction of the plant Aglaophyton major by D. S. Edwards (1986). The original name, given by Kidson and Lang, was Rhynia major, but Edwards showed that the central strand did not consist of tracheids with secondary wall thickenings. So the plant had to be transferred to another genus. In fact Aglaophyton cannot be reckoned to belong to the Tracheophyta and does have affinities to the mosses. Aglaophyton major may represent an intermediate stage in the evolution of vascular plant known as protracheophytes. The transverse section of the stem resembles that of Rhynia gwynne-vaughanii, but it is larger. The size of the plant is estimated at a maximum of 18 cms. The stem diameter is between 1.5 and 6 mms.

Phylum Zosterophyllophyta
Fossils of zosterophylls have been found in rocks from the Early and Middle Devonian, 408 to 370 million years ago.

  • Aerial stems were covered with cuticle and had stomata only on the upper branches.

  • Zosterophyllum dichotomously branched, but frequently lateral branches further differentiated into one axis that grew upward and another downward.

  • The lower branch may have function as a root.

  • Some Zosterophylls were naked or had small spine-like enations.

  • The sporangia were globose or reniform and borne laterally on short stalks.

  • Zosterophylls were homosporous.

  • Xylem matured from the periphery toward the center: centripetal differentiation.

  • They are considered to be the ancestral group that gave rise to the Lycopods; the sporangia of both groups are borne laterally and are similar in shape, and the xylem in both groups differentiates centripetally.

Phylum Trimerophytophyta
This phylum probably evolved directly from the rhyniophytes.
Trimerophytes were larger and more complex plants than the rhyniophytes or zosterophyllophytes.
Trimerophytes appeared in the Early Devonian about 395 million years ago and had become extinct by the end of the mid Devonian, about 20 million years later – a relative short period of existence.

  • They lacked leaves.

  • Lateral branches forked dichotomously several times.

  • They were homosporous.

  • Some branches were vegetative while others bore elongated sporangia.

  • Vascular strand was more massive than that of the rhyniophytes.

  • The xylem differentiated centrifugally.

Phylum Lycopodiophyta
There are about 15 genera of lycophytes and approximately 1200 living species.
This evolutionary line extends back into the Devonian but was most prevalent in the wet swamps of the Carboniferous period.
Molecular and morphological evidence indicate that they split up into two evolutionary lines before 400 million years ago, in the Early – Mid Devonian.

  • The lycophyte clade includes the modern lycophytes.

  • The euphyllophytes clade includes whisk ferns, horsetails and seed plants.

The extinct lycophytes include very large woody trees that did not survive in the drier climate at the end of and after the Carboniferous age. In the Carboniferous some lycophytes were forest-forming trees more than 35 meters tall.

Woody lycophytes became extinct before the end of the Paleozoic era, 248 million years ago.
The second and the surviving group of Lycopods are the small and herbaceous plants.
Lycophyta remains became the largest coal deposits of all geologic time.
Lycophyta are characterized by

  • microphyllous leaves,

  • a special spore producing body called a strobilus,

  • the presence of true vascular stems, roots and leaves.

There are three prominent orders of Lycophyta:

  • Lycopodiales, or club mosses;

  • Selaginellales, or Spike mosses;

  • Isoetales, or Quillworts.

Family Lycopodiaceae
A family of about 400 species, mostly tropical.
The taxonomic boundaries of the genera are not well understood and as many as 15 genera may eventually be recognized. Seven of these genera are represented in North America.

  • Sporophyte with true leaves, stems and roots.

  • Dichotomous branching rhizome from which aerial branches and roots arise.

  • Stems and roots are protostelic or siphonostelic. Leaf gaps absent.

  • Leaves are microphyllous and spirally arranged, sometimes opposite or whorled.

  • Sporophylls, modified fertile microphylls, sometimes grouped into strobili (strobilus, cluster of overlapping non-photosynthetic sporophylls).

  • In Huperzia and Phlegmariurus, the sporophylls are similar to ordinary microphylls and are interspersed among the sterile microphylls.

  • One sporangium per sporophyll, near the base and on the adaxial side.

  • Homosporous.

  • Gametangia, antheridia and archegonia, may require 6 to 15 years to mature.

  • Self-fertilization is rare.

  • Biflagellated sperm requires water to reach the archegonium.

Family Selaginellaceae
There are about 700 species in this family, most of them tropical. Selaginella is the only genus in the family.

  • Plants herbaceous, annual or perennial, sometimes remaining green over winter.

  • Stems leafy, branching dichotomously, regularly or irregularly forked or branched.

  • Stems and roots protostelic (sometimes with many protosteles or meristeles), siphonostelic, or actino-plectostelic.

  • Protostele held in place by trabeculae.

  • Rhizophores (modified leafless shoots producing roots) present or absent, geotropic, borne on stems at branch forks, throughout, or confined to base of stems.

  • Leaves microphylls, on 1 plant dimorphic or monomorphic, small, with adaxial ligule near base, single-veined, rarely veins forked.

  • Strobili sometimes ill-defined, terminal, cylindrical, quadrangular, or flattened.

  • Sporophylls (fertile leaves) monomorphic or adjacently different, slightly or highly differentiated from vegetative (sterile) leaves: microsoporophylls and megasporophylls.

  • Sporophylls and microphylls with ligule.

  • Sporangia short-stalked, solitary in axil of sporophylls, opening by distal slits.

  • Spores of 2 types (plants heterosporous), megaspores (1--2--4), large; microspores numerous (hundreds), very small.

  • Gametophytes unisexual.

  • Gametophytes develop inside the spore wall: endosporic development.

  • Microgametophyte lacks chlorophyll.

  • At maturity, it consists of a single prothallial cell and an antheridium.

  • Antheridium produces many of biflagellated sperms.

  • Microspore wall ruptures to liberate the sperms.

  • Megagametophyte multicellular.

  • Megagametophyte protrudes through a rupture in the spore wall.

  • Archegonia develop in the exposed area.

  • Sperms require water to swim to the archegonia.

  • The suspensor develops and pushes the developing embryo deep into the female gametophyte.

Family Isoetaceae
A family of one genus, Isoetes (ca. 150 spp.), found world-wide, especially in temperate areas.
A second genus, Stylites, is sometimes recognized.
Some authors put fossil representatives in the genus Isoetites, which is known from as early as the upper Triassic.

  • Plants tufted, grass-like, perennial, evergreen aquatics to ephemeral terrestrials.

  • Underground stem brown, cormlike, lobed.

  • Roots arising along central groove separating each rootstock lobe, simple or dichotomously branched, containing eccentric vascular strand and surrounding lacuna.

  • Leaves linear, simple, spirally or distichously arranged, dilated toward base, tapering to apex, containing 4 transversely septate longitudinal lacunae, a central collateral vascular strand, and frequently several peripheral fibrous bundles.

  • Each leaf is a potential sporophyll.

  • Ligule inserted above sporangium on each sporophyll.

  • Heterosporous (megaspores and microspores not alike).

  • Megasporophylls and microsporophylls usually borne in alternating cycles; hardened scales and phyllopodia occasionally surround leaves.

  • Sporangia solitary, adaxial, embedded in basal cavity of leaf, velum (thin flap extending downward over sporangium) partly to completely covering adaxial surface of sporangium.

  • Megasporangium with several to hundreds of megaspores.

  • Microsporangium with thousands of microspores.

  • Megagametophytes white, endosporic, exposed when megaspore opens along proximal ridges; archegonia 1 to several, indicated by quartets of brownish neck cells.

  • Microgametophytes 9-celled, endosporic, antheridium releasing 4 multitailed spermatozoids.

Phylum Trimerophytophyta.
A fossil phylum that seems to have evolved from rhyniophytes.
It might be the ancestor of ferns, progymnosperms and perhaps horsetails.
They first appeared in the early Devonian about 395 m.y.a. and became extinct by the end of the mid Devonian, at about 370 m.y.a.

  • Trimerophytes lacked leaves and roots; most of the plant body consisted of branching stems that were photosynthetic throughout their length.

  • Vascular tissue was present, forming a solid central bundle in the center of the stem, or protostele.

  • Their protostele was more massive than that of rhyniophytes.

  • They had a band of thick-cell-wall cells in the cortex.

  • Xylem differentiated centrifugally like in rhyniophytes.

  • Trimerophytes branched pseudomonopodially, that means that the branching was unequal, forming a main stem, or axis, with several smaller lateral branches.

  • Rhyniophytes branched dichotomously: stems always branched into two equal branches.

  • Lateral branches typically branched dichotomously, and were often shortened to form bushy "webs" of small, closely spaced branches.

  • Some trimerophytes also bore enations on the main stems, giving them a superficially "thorny"


  • Trimerophytes bore sporangia at the tips of branches, like the rhyniophytes, but unlike the superficially similar zosterophylls from the same time period,.

  • The spindle-shaped sporangia produced only one type of spore: trimerophytes were homosporous.

  • Spores released from the sporangia would have germinated into gametophytes, but no fossil trimerophytes gametophytes have been identified, and we do not know whether trimerophytes, like their later relatives the ferns, sphenopsids, and seed plants, had small, inconspicuous gametophytes.

  • Trimerophytes varied in size from a few centimeters to nearly a meter tall; large trimerophytes were among the largest plants of the Early Devonian.

Phylum Psilophyta
This phylum is also known as Psilotophyta.
It includes two living genera, Psilotum and Tmesipteris, from tropical and subtropical regions of the world..

  • Sporophyte with a dichotomously branching aerial and subterranean stem system.

  • True roots lacking.

  • Aerial stems lacking leaves but with scale-like or larger leaf-like structures (enations)

  • Gametophytes bisexual, subterranean, resembling a small piece of rhizome

  • Gametophyte with a symbiotic fungus. Some have a vascular tissue

  • The sperm is multiflagellated and requires water to swim to the archegonium.

  • Homosporous

  • Sporangia, a 2 or 3-chambered synangium, borne at the apex of small side-branches (appear to be arranged along the sides of the major stems of the plant--sporophyte)

  • Sporophyte remains attached to the gametophyte in the early stages by a foot and derives nourishment from the gametophyte.

Phylum Sphenopsida
Sphenopsids extend back to the Devonian and reached their maximum development in the Carboniferous (380-280 m.y.a.).
A family of one extant genus, Equisetum (ca. 15 species), of nearly worldwide distribution in damp habitats such as riverbanks, lakeshores, and marshes. Michigan is a center of diversity for the genus with nine native species.

  • The sporophyte of Equisetum is differentiated into an underground rhizome that bears adventitious roots and an upright, photosynthetic stem with whorls of microphylls..

  • Tough perennial herbs with jointed, ridged aerial stems with distinct nodes.

  • The aerial stems contain a large central pith region, which in mature plants is hollow.

  • Surrounding the pith cavity are discrete bundles of vascular tissue; this arrangement of conducting tissue is known as a eustele.

  • The bundles contain both xylem and phloem, and are marked by the presence of large canals known as carinal canals (under the ridges), which also function in water conduction.

  • External to the vascular bundles is another set of canals, the vallecular canals or cortical canals (under the valleys).

  • These canals line up with the depressions between the ribs on the surface of the stem. Most fossil sphenophytes had very similar stem morphology.

  • Leaves are small, whorled, non-photosynthetic, fused together to various degrees and adpressed to the stem.

  • Sporangia borne on peltate sphorangiophores aggregated into a strobilus.

  • Strobili consist of tightly packed appendages called sporangiophores.

  • Homosporous; spores with elaters.

  • Gametophytes green, epigeal, bisexual or unisexual, male gametophytes smaller than female; with rhizoids.

  • Archegonia develop before antheridia to increase the probability of cross-fertilization.

  • Sperms are multiflagellated and require water to swim to the egg.

  • Several eggs on a single gametophyte may be fertilized.

Phylum Pterophyta
The fossil record of ferns extends back into the Carboniferous
There are 11,000 species of ferns in the world. Most species are tropical.

  • There are about 380 species of ferns in U.S.A. and Canada.

  • Costa Rica has over 1,000 species.

  • Philippines has over 950 species.

  • Sporophyte is differentiated into true roots, stem (rhizome) and leaves (megaphylls).

  • The xylem is composed of tracheids.

  • Leaves or fronds develop in curled position at the apex of the rhizome (stem) and uncoil as they mature, a condition called circinate vernation.

  • Leaves usually differentiated into stipe (petiole) and blade with a central rachis or vein.

  • Leaf gaps are conspicuous.

  • Most ferns are homosporous; a few aquatic genera are heterosporous.

  • Sporangia normally develop on the abaxial (dorsal, lower side) or on the margin of the leaf.

  • Gametophytes are inconspicuous and arise directly from spores.

  • Gametophytes are bisexual in homosporous ferns and unisexual in heterosporous ferns.

  • Sperms are flagellated and coiled.

Sporangial development is important in the understanding of the evolutionary relationship of vascular plants.

  1. The eusporangiate sporangium (eusporangia) has the sporangial wall developing from several superficial cells and the sporogenous tissue develops from internal cells of the sporophyll. The sporangial wall is several cells thick. The inner wall cells may be compressed and stretched so that the sporangial wall may appear to be made of a single layer of cells at maturity. The sporangia are large and contain a large number of spores.

  1. The leptosporangiate sporangium (leptosporangia) develops entirely from the periclinal division of a single superficial cell. By a precise pattern of division a stalk and a globose capsule are formed. A nutritive tapetum develops inside the capsule. The inner mass of cells develops into spore mother cells that undergo meiosis. The remnant of the tapetum is deposited around the spores and gives them their characteristic ornamentation. The annulus is made of unevenly thickened cells. The number of spores produced is a multiple of 2 (between 16 and 512 in homosporous species).

  1. Eusporangiate sporangium is characteristic of all vascular plants except the leptosporangiate ferns.

There are five orders of ferns often recognized by pteridologists.

  • Ophioglossales, Marattiales, Filicales, Marsileales and Salviniales.

The Ophioglossales and Marattiales are eusporangiate. The other three are leptosporangiate.

Order Ophioglossales.
There are three genera in the Ophioglossales.

  • Ophioglossales are eusporangiate.

  • Two genera, Botrychium and Ophioglossum, are widespread in the world.

  • A single leaf is produced each year.

  • Each leaf consists of two parts: a vegetative portion or blade and a fertile segment.

  • The vegetative portion is dissected in Botrychium and undivided in Ophioglossum.

  • Their gametophyte is subterranean, with many rhizoids and associated to an endophytic fungus.

Order Marattiales.
This is an ancient group of eusporangiate ferns with fossil members extending back into the Carboniferous.

  • They are large ferns with dissected fronds.

  • There are six living genera with about 200 species.

Order Filicales.
There are about 35 families, 320 genera and 10,500 species in this order.

  • The Filicales are leptosporangiate, which differentiate them form Ophioglossales and Marattiales.

  • The Filicales are homosporous while the Marsileales and Salviniales are heterosporous.

  • Megaphylls vary from entire to highly dissected (pinnate).

  • Stipe, rachis, pinnae, pinnules.

  • Uncoiling of young leaves is influenced by auxin, a plant hormone.

  • Scales called paleae, and hairs of different kinds often cover the rhizome and young fronds.

  • Sporangia may cover the dorsal surface of the blade or may be grouped into sori (sorus).

  • A sorus may or may not be protected by an indusium.

  • Spores give rise to free living bisexual gametophytes called a prothallus.

  • The gametophyte is chlorophyllous, heart-shaped, with rhizoids.

  • Gametangia develop on the underside of the prothallus.

  • Water is required for fertilization to occur.

  • In some ferns from different parts of the world produce gametophytes that persist indefinitely without forming sporophytes.

  • These gametophytes reproduce asexually by outgrowths called gemmae.

Order Marsileales and Salviniales.

  • Both orders are heterosporous leptosporangiate ferns.

  • The sporophyte grows in mud or submerged with leaves floating on the water surface.

  • Marsileales produce bean-shaped drought resistant sporocarps.

  • Sporocarps germinate to produce a chain of sori each bearing megasporangia and microsporangia.

  • Their gametophytes are very specialized.

  • There are three genera of Marsileales and about 72 species.

  • There are two general of Salviniales, Salvinia and Azolla.

  • These are small, floating plants.

  • Azolla produces their spores in sporocarps.

  • Azolla leaves are modified to accommodate filaments of Anabaena azollae, nitrogen-fixing cyanobacteria.

  • Salvinia produces a whorl of three leaves, tow of which are chlorophyllous and undivided, and the third is divided and hags down the surface of the water resembling a root.

  • Sterile leaves have water repellent hairs.

  • The submerged leaf produces a sporocarp with a single sorus.

  • Gametophytes are very small.

  • Megagametophyte produces a single archegonium.

  • Microgametophytes germinate inside the sporocarp and produce two antheridia.

  • Each antheridium releases four motile sperms.

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