BSC 1011C
General Biology II
Dr. Graeme Lindbeck
glindbeck@valenciacollege.edu

Plant Diversity II: The Evolution of Seed Plants

Outline

Introduction

The evolution of plants is highlighted by two important landmarks:

the evolution of seeds, which lead to the gymnosperms and angiosperms, the plants that dominate most modern landscapes
the emergence of the importance of seed plants to animals, specifically to humans.

Agriculture, the cultivation and harvest of plants (primarily seed plants), began approximately 10,000 years ago in Asia, Europe, and the Americas.

This was the single most important cultural change in the history of humanity, for it made possible the transition from hunter-gather societies to permanent settlements.

The seeds and other adaptations of gymnosperms and angiosperms enhanced the ability of plants to survive and reproduce in diverse terrestrial environments.

Plants became the main producers on land.

Seed plants are vascular plants that produce seeds.

Contributing to the success of seed plants as terrestrial organisms are three important reproductive adaptations:

continued reduction of the gametophyte
the advent of the seed
the evolution of pollen.

A. Overview of Seed Plant Evolution

1. Reduction of the gametophyte continued with the evolution of seed plants

An important distinction between mosses and other bryophytes and ferns and other seedless vascular plants is a gametophyte-dominated life cycle for bryophytes and a sporophyte-dominant life cycle for seedless vascular plants.

Continuing that trend, the gametophytes of seed plants are even more reduced than those of seedless vascular plants such as ferns.

In seeds plants, the delicate female gametophyte and young embryos are protected from many environmental stresses because they are retained within the moist sporangia of the parental sporophyte.

The gametophytes of seed plants obtain nutrients from their parents, while those of seedless vascular plants are free-living and fend for themselves.

For the gametophyte to exist within the sporophyte has required extreme miniaturization of the the gametophyte of seed plants.

The gametophytes of seedless vascular plants are small but visible to the unaided eye, while those of seed plants are microscopic.

Why has the gametophyte generation not been completely eliminated from the plant life cycle?

The haploid generation may provide a mechanism for "screening" new alleles, including mutations.
- Gametophytes with deleterious mutations affecting metabolism or cell division will not survive to produce gametes that could combine to start new sporophytes.
Another possible reason is that all sporophyte embryos are dependent, at least to some extent, on tissues of the maternal gametophyte.
- The gametophyte nourishes the sporophyte embryo, at least during its early development.

2. Seeds became an important means of dispersing offspring

In bryophytes and seedless vascular plants, spores from the sporophyte are the resistant stage in the life cycle.

For example, moss spores can survive even if the local environment is too extreme for the moss plants themselves to survive.
Because of their tiny size, the spores themselves might also be dispersed in a dormant state to a new area.

Spores were the main way that plants spread over Earth for the first 200 millions years of life on land.

The seed represents a different solution to resisting harsh environments and dispersing offspring.

In contrast to a single-celled spore, a multicellular seed is a more complex, resistant structure.

A seed consists of a sporophyte embryo packaged along with a food supply within a protective coat.

There are evolutionary and developmental relationships between spores and seeds.

The parent sporophyte does not release its spores, but retains them within its sporangia.
Not only are the spores retained, but the gametophyte develops within the spore from which it is derived.

All seed plants are heterosporous, producing two different types of sporangia that produce two types of spores.

Megasporangia produce megaspores, which give rise to female (egg-containing) gametophytes.
- Microsporangia produce microspores, which give rise to male (sperm-containing) gametophytes.

In contrast to heterosporous seedless vascular plants, the megaspores and the female gametophytes of seed plants are retained by the parent sporophyte.

Layers of sporophyte tissues, integuments, envelop and protect the megasporangium.

An ovule consists of integuments, megaspore, and megasporangium.

A female gametophyte develops inside a megaspore and produces one or more egg cells.
A fertilized egg develops into a sporophyte embryo.
The whole ovule develops into a seed.

A seed's protective coat is derived from the integuments of the ovule.

Within this seed coat, a seed may remain dormant for days, months, or even years until favorable conditions trigger germination.

When the seed is eventually released from the parent plant, it may be close to the parent, or be carried off by wind or animals.

3. Pollen eliminated the liquid-water requirement for fertilization

The microspores, released from the microsporangium, develop into pollen grains.

These are covered with a tough coat containing sporopollenin.

They are carried away by wind or animals until pollination occurs when they land in the vicinity of an ovule.

The pollen grain will elongate a tube into the ovule and deliver one or two sperm into the female gametophyte.

While some primitive gymnosperms have flagellated sperm cells, the sperm in most gymnosperms and all angiosperms lack flagella.

In seed plants, the use of resistant, far-traveling, airborne pollen to bring gametes together is a terrestrial adaptation.

In bryophytes and pteridophytes, flagellated sperm must swim through a film of water to reach eggs cells in archegonia.

The evolution of pollen in seed plants led to even greater success and diversity of plants on land.

4. The two clades of seed plants are gymnosperms and angiosperms

Like other groups of organisms, our understanding of plant taxonomy is being revised to reflect new data, new methods, and new ideas.

The current data support a phylogeny of the seed plants with two main monophyletic branches - the gymnosperms and the angiosperms.

Both probably evolved from different ancestors in an extinct group of plants, the progymnosperms, some of which had seeds.

B. Gymnosperms

The most familiar gymnosperms are the conifers, the cone-bearing plants such as pines.

The ovules and seeds of gymnosperms ("naked seeds") develop on the surfaces of specialized leaves called sporophylls.

In contrast, ovules and seeds of angiosperms develop in enclosed chambers (ovaries).

Gymnosperms appears in the fossil record much earlier than angiosperms.

1. The Mesozoic era was the age of gymnosperms

The gymnosperms probably descended from progymnosperms, a group of Devonian plants.

While the earliest progymnosperms lacked seeds, by the end of the Devonian, some species had evolved seeds.

Adaptive radiation during the Carboniferous and early Permian produced the various phyla of gymnosperms.

The flora and fauna of Earth changed dramatically during the formation of the supercontinent Pangaea in the Permian.

This likely led to major environmental changes, including drier and warmer continental interiors.

Many groups of organisms disappeared and others emerged as their successors.

For example, amphibians decreased in diversity while reptiles increased.
Similarly, the lycophytes, horsetails, and ferns that dominated in Carboniferous swamps were largely replaced by gymnosperms, which were more suited to the drier climate.

The change in organisms was so dramatic that geologist use the end of the Permian, about 245 million years ago, as the boundary between the Paleozoic and Mesozoic eras.

The terrestrial animals of the Mesozoic, including dinosaurs, were supported by a vegetation consisting mostly of conifers and cycads, both gymnosperms.

The dinosaurs did not survive the environmental upheavals at the end of the Mesozoic, but many gymnosperms persisted and are still an important part of Earth's flora.

2. The four phyla of extant gymnosperms are ginko, cycads, gnetophytes, and conifers

There are four plant phyla grouped as gymnosperms.

Phylum Ginkgophyta consists of only a single extant species, Ginkgo biloba.

This popular ornamental species has fanlike leaves that turn gold before they fall off in the autumn.
Landscapers usually only plant male trees because the seed coats on female plants decay, they produce a repulsive odor (to humans, at least).

Cycads (phylum Cycadophyta) superficially resemble palms.

Palms are actually flowering plants.

Phylum Gnetophyta consists of three very different genera.

Weltwitschia plants, from deserts in southwestern Africa, have straplike leaves.
Gentum species are tropical trees or vines.
Ephedra (Mormon tea) is a shrub of the American deserts.

3. The life cycle of a pine demonstrates the key reproductive adaptations of seed plants

The life cycle of a pine illustrates the three key adaptations to terrestrial life in seed plants:

increasing dominance of the sporophyte
seeds as a resistant, dispersal stage
pollen as an airborne agent bringing gametes together.

The pine tree, a sporophyte, produces its sporangia on scalelike sporophylls that are packed densely on cones.

Conifers, like all seed plants, are heterosporous, developing male and female gametophytes from different types of spores produced by separate cones.

Each tree usually has both types of cones.
Small pollen cones produce microspores that develop into male gametophytes, or pollen grains.
Larger ovulate cones make megaspores that develop into female gametophytes.

It takes three years from the appearance of young cones on a pine tree to the formation mature seeds.

The seeds are typically dispersed by the wind.

Reproduction in pines begins with the appearance of cones on a pine tree.

Most species produce both pollen cones and ovulate cones.
A pollen cone contains hundreds of microsporangia held on small sporophylls.
- Cells in the microsporangia undergo meiosis to form haploid microspores that develop into pollen grains.
An ovulate cone consists of many scales, each with two ovules.
- Each ovule includes a megasporangium.
During pollination, windblown pollen falls on the ovulate cone and is drawn into the ovule through the micropyle.
- The pollen grain germinates in the ovule, forming a pollen tube that digests its way through the megasporangium.
The megaspore mother cell undergoes meiosis to produce four haploid cells, one of which will develop into a megaspore.
- The megaspore grows and divides mitotically to form the immature female gametophyte.
Two or three archegonia, each with an egg, then develop within the gametophyte.
At the same time that the eggs are ready, two sperm cells have developed in the pollen tube which has reached the female gametophyte.
- Fertilization occurs when one of the sperm nuclei fuses with the egg nucleus
The pine embryo, the new sporophyte, has a rudimentary root and several embryonic leaves.
- The female gametophyte surrounds and nourishes the embryo.
- The ovule develops into a pine seed, which consists of an embryo (new sporophyte), its food supply (derived from gametophyte tissue), and a seed coat derived from the integuments of the parent tree (parent sporophyte).

The conifers, phylum Coniferophyta, is the largest gymnosperm phylum.

The term conifer comes from the reproductive structure, the cone, which is a cluster of scalelike sporophylls.
Although there are only about 550 species of conifers, a few species dominate vast forested regions in the Northern Hemisphere where the growing season is short.

Conifers include pines, firs, spruces, larches, yews, junipers, cedars, cypresses, and redwoods.

Most conifers are evergreen, retaining their leaves and photosynthesizing throughout the year.

Some conifers, like the dawn redwood and tamarack, are deciduous, dropping their leaves in autumn.

The needle-shaped leaves of some conifers, such as pines and firs, are adapted for dry conditions.

A thick cuticle covering the leaf and the placement of stomata in pits further reduce water loss.

Much of our lumber and paper comes from the wood (actually xylem tissue) of conifers.

This tissue gives the tree structural support.

Coniferous trees are amongst the largest and oldest organisms of Earth.

Redwoods from northern California can grow to heights of over 100m.
One bristlecone pine, also from California, is more than 4,600 years old.

C. Angiosperms (Flowering Plants)

Angiosperms, better known as flowering plants, are vascular seed plants that produce flowers and fruits.

They are by far the most diverse and geographically widespread of all plants.

There are abut 250,000 known species of angiosperms.

1. Systematists are identifying the angiosperm clades

All angiosperms are placed in a single phylum, the phylum Anthophyta.

As late as the 1990s, most plant taxonomists divided the angiosperms into two main classes, the monocots and the dicots.

Most monocots have leaves with parallel veins, while most dicots have netlike venation.

Recent systematic analyses have upheld the monocots as a monophyletic group.

They include lilies, orchids, yuccas, grasses, and grains.

However, molecular systematics has indicated that plants with the dicot anatomy do not form a monophyletic group.

One clade, the eudicots, does include the majority of dicots.

It includes roses, peas, sunflowers, oaks, and maples.

Some other dicots actually belong to angiosperm lineages that diverged earlier that the origin of either monocots or eudicots.

These include the star anise, the water lilies, and Amborella trichopoda from the oldest angiosperm branch.

While most angiosperms belong to either the monocots (65,000 species) or eudicots (165,000 species) several other clades branched off before these.

Based on molecular analyses, Arborella is the only survivor of a branch at the base of the angiosperm tree.

Refinements in vascular tissue, especially xylem, probably played a role in the enormous success of angiosperms in diverse terrestrial habitats.

Like gymnosperms, angiosperms have long, tapered tracheids that function for support and water transport.
Angiosperms also have fibers cells, specialized for support, and vessel elements (in most angiosperms) that develop into xylem vessels for efficient water transport.

2. The flower is the defining reproductive adaptation of angiosperms

While evolutionary refinements of the vascular system contributed to the success of angiosperms, the reproductive adaptations associated with flowers and fruits contributed the most.

The flower is an angiosperm structure specialized for reproduction.

In many species, insects and other animals transfer pollen from one flower to female sex organs of another.
Some species that occur in dense populations, like grasses, rely on the more random mechanism of wind pollination.

A flower is a specialized shoot with four circles of modified leaves: sepals, petals, stamens, and carpals.

The sepals at the base of the flower are modified leaves that enclose the flower before it opens.

The petals lie inside the ring of sepals.

These are often brightly colored in plant species that are pollinated by animals.
They typically lack bright coloration in wind-pollinated plant species.

Neither the sepals or petals are directly involved in reproduction.

Stamens, the male reproductive organs, are the sporophylls that produce microspores that will give rise to gametophytes.

A stamen consists of a stalk (the filament) and a terminal sac (the anther) where pollen is produced.

Carpals are female sporophylls that produce megaspores and their products, female gametophytes.

At the tip of the carpal is a sticky stigma that receives pollen.
A style leads to the ovary at the base of the carpal.
Ovules and, later, seeds are protected within the ovary.

The enclosure of seed within the ovary (the carpal), a distinguishing feature of angiosperms, probably evolved from a seed-bearing leaf that became rolled into a tube.

Some angiosperms have flowers with single carpals (garden peas), others have several separate carpals (magnolias) or fused carpals (lilies).

3. Fruits help disperse the seeds of angiosperms

A fruit is a mature ovary.

As seeds develop from ovules after fertilization, the wall of the ovary thickens to form the fruit.
Fruits protect dormant seeds and aid in their dispersal.

Various modifications in fruits help disperse seeds.

In some plants, such as dandelions and maples, the fruit functions like a kite or propeller, enhancing wind dispersal.

Many angiosperms use animals to carry seeds.

Fruits may be modified as burrs that cling to animal fur.
Edible fruits are eaten by animals when ripe and the seeds are deposited unharmed, along with fertilizer.

The fruit develops after pollination triggers hormonal changes that cause ovarian growth.

The wall of the ovary becomes the pericarp, the thickened wall of the fruit.
The other parts of the flower whither away in many plants.
If a flower has not been pollinated, the fruit usually does not develop, and the entire flower withers and falls away.

Fruits are classified into several types depending on their developmental origin.

Simple fruits are derived from a single ovary.
- These may be fleshy, such as a cherry, or dry, such as a soybean pod.
An aggregate fruit, such as a blackberry, results from a single flower with several carpals.
A multiple fruit, such as a pineapple, develops from an inflorescence, a tightly clustered group of flowers.

By selectively breeding plants, humans have capitalized on the production of edible fruits.

Apples, oranges, and other fruits in grocery stores are exaggerated versions of much smaller natural varieties of fleshy fruits.

The staple foods for humans are the dry, wind-dispersed fruits of grasses.

These are harvested while still on the parent plant.
The cereal grains of wheat, rice, corn, and other grasses are actually fruits with a dry pericarp that adheres tightly to the seed coat of the single seed inside.

4. The life cycle of an angiosperm is a highly refined version of the alternation of generations common in plants

All angiosperms are heterosporous, producing microspores that form male gametophytes and megaspores that form female gametophytes.

The immature male gametophytes are contained within pollen grains and develop within the anthers of stamens.
- Each pollen grain has two haploid cells.
Ovules, which develop in the ovary, contain the female gametophyte, the embryo sac.
- It consists of only a few cells, one of which is the egg.

The life cycle of an angiosperm begins with the formation of a mature flower on a sporophyte plant and culminates in a germinating seed.

The anthers of the flower produce microspores that form male gametophytes (pollen).
Ovules produce megaspores that form female gametophytes (embryo sacs).
After its release from the anther, pollen is carried to the sticky stigma of a carpal.
- Although some flowers self-pollinate, most have mechanisms that ensure cross-pollination, transferring pollen from flowers of one plant to flowers of another plant of the same species.
- The pollen grain germinates (begins growing) from the stigma toward the ovary.
- When the pollen tube reaches the micropyle, a pore in the integuments of the ovule, it discharges two sperm cells into the female gametophyte.
In a process known as double fertilization, one sperm unites with the egg to form a diploid zygote and the other fuses with two nuclei in the large center cell of the female gametophyte.
The zygote develops into a sporophyte embryo packaged with food and surrounded by a seed coat.
- The embryo has a rudimentary root and one or two seed leaves, the cotyledons.
- Monocots have one seed leaf and dicots have two.

Monocots store most of the food for the developing embryo in endosperm which develops as a triploid tissue in the center of the embryo sac

Beans and many dicots transfer most of the nutrients from the endosperm to the developing cotyledons.

One hypothesis for the function of double fertilization is that it synchronizes the development of food storage in the seed with development of the embryo.

Double fertilization may prevent flowers from squandering nutrients on infertile ovules.

The seed consists of the embryo, endosperm, sporangium, and a seed coat from the integuments.

As the ovules develop into seeds, the ovary develops into a fruit.

After dispersal by wind or animals, a seed germinates if environmental conditions are favorable.

During germination, the seed coat ruptures and the embryo emerges as a seedling.
It initially uses the food stored in the endosperm and cotyledons to support development.

5. The radiation of angiosperms marks the transition from the Mesozoic era to the Cenozoic era

Earth's landscape changed dramatically with the origin and radiation of flowering plants.

The oldest angiosperm fossils are found in rocks in the early Cretaceous, about 130 million years ago.

By the end of the Cretaceous, 65 million years ago, angiosperms had become the dominant plants on Earth.

6. Angiosperms and animals have shaped one another's evolution

Ever since they colonized the land, animals have influenced the evolution of terrestrial plants and vice versa.

The fact that animals must eat affects the natural selection of both animals and plants.

Natural selection must have favored plants that kept their spores and gametophytes far above the ground, rather than dropping them within the reach of hungry ground animals.
In turn, this may have been a selective factor in the evolution of flying insects.

On the other hand, some herbivores may have become beneficial to plants by carrying the pollen and seeds of plants that they used as food.

Natural selection reinforced these interactions, for they improved the reproductive success of both partners.

This type of mutual evolutionary influence between two species is termed coevolution.

Pollinator-plant relationships are partly responsible for the diversity of flowers.

In many cases, a plant species may be pollinated by a group of pollinators, such as diverse species of bees or hummingbirds, and have evolved flower color, fragrance, and structures to facilitate this.
Conversely, a single species, such as a honeybee species, may pollinate many plant species.

D. Plants and Human Welfare

The absolute dependence of humans on Earth's flora is a specific and highly refined case of the more general connection between animals and plants.

Like other organisms, we depend on photosynthetic organisms for food production and oxygen release.

However, we use technology to manipulate or select plants that maximize the harvest of plant products for human use.

1. Agriculture is based almost entirely on angiosperms

Flowering plants provide nearly all our food.

All of our fruit and vegetable crops are angiosperms.
Corn, rice, wheat, and other grain are grass fruits.
- The endosperm of the grain seeds is the main food source for most of the people of the world and their domesticated animals.

We also grow angiosperms for fiber, medications, perfumes, and decoration.

Like other animals, early humans probably collected wild seeds and fruits.

Agriculture developed gradually as humans began sowing seed and cultivating some plant species to provide a more dependable food source.

As they domesticated certain plants, they used selective breeding to improve the quantity and quality of the foods the crops produced.

2. Plant diversity is a nonrenewable resource

The demand for space and natural resources resulting from the exploding human population is extinguishing plant species at an unprecedented rate.

This is especially acute in the tropics where half the human population lives and where growth rates are highest.

Due primarily to the slash-and-burn clearing of forests for agriculture, tropical forests may be completely eliminated within 25 years.

As the forests disappear, thousand of plants species and the animals that depend on these plants also go extinct.

The destruction of these areas is an irrevocable loss of these nonrenewable resources.

While the loss of species is greatest in the tropics, this environmental assault occurs worldwide.

In addition to the ethical concerns that many people have concerning the extinction of living forms, there are also practical reasons to be concerned about the loss of plant diversity.

We depend on plants for food, building materials, and medicines.

We have explored the potential uses for only a tiny fraction of the 250,000 known plant species.
Almost all of our food is based on cultivation of only about two dozen species.

We have derived many medical compounds from the unique secondary compounds of plants.

More than 25% of prescription drugs are extracted from plants, and many more medicinal compounds were first discovered in plants and then synthesized artificially.

Researchers have investigates fewer than 5,000 plant species as potential sources of medicines.

Pharmaceutical companies were led to most of these species by local people who use the plants in preparing their traditional medicines.

The tropical rain forests and other plant communities may be a medicine chest of healing plants that could be extinct before we even know they exist.

We need to view rain forests and other ecosystems as living treasures that we can harvest only at sustainable rates.

Course Pages maintained by
Dr. Graeme Lindbeck .