Evolution of Plants

Modern classification systems, based largely on molecular evidence, divide living organisms into three domains:
Bacteria (also called Eubacteria), Archaea, and Eukarya. Plants are classified as a kingdom (Plantae) within the
Eukarya; organisms that possess a nucleus, mitochondria, an internal cytoskeleton , and, in photosynthetic species,
chloroplasts. Most scientists recognize three other eukaryotic kingdoms: Protista (most of which are single-celled
organisms), Fungi, and Animalia (animals). The fungi, plants, and animals are thought to have evolved from different
groups of protists.
Plants are multicellular organisms that have evolved the ability to live on land. The vast majority can carry out
photosynthesis, but they are not the only organisms with this ability: many protists can photosynthesize too, as can
several important groups of bacteria.
Algae in Plant Evolution
Photosynthetic protists (commonly called algae) are a diverse group of organisms and are divided into several phyla.
Many are unicellular, including most euglenoids (phylum Euglenophyta) and dinoflagellates (Dinophyta), and some
diatoms (Bacillariophyta) and green algae (Chlorophyta). These, along with the cyanobacteria (often misleadingly
called blue-green algae), form the phytoplankton of aquatic ecosystems. Others, including all brown algae
(Phaeophyta), most red algae (Rhodophyta), and many green algae are multicellular. The large marine forms of these
phyla are usually called seaweeds.
Plants are thought to have evolved from a class of freshwater green algae called the charophytes. Two groups of
charophyte, the Coleochaetales and the Charales, resemble the earliest land plants (bryophytes) in a variety of ways,
including the structure of their chloroplasts and sperm cells, and the way their cells divide during mitosis.
The Importance of Vascular Tissue
Plants are classified into two main groups: the bryophytes (nonvascular plants) and the tracheophytes (vascular plants).
Both groups have multicellular embryos, which indicates that they are closely related to each another and distinguishes
them from the green algae. Indeed, true plants are often referred to as embryophytes because of this feature. The
bryophytes consist of the liverworts, hornworts, and mosses, and as their name implies none of these plants possess
vascular tissues.
All other plants, including the ferns, gymnosperms, and angiosperms, are classified as tracheophytes. These possess
specialized vascular tissues—phloem and xylem —to transport sugars, water, and minerals throughout their bodies.
The oldest known vascular plants appeared in the middle Silurian period (439–409 million years ago); the oldest known
bryophytes appeared later, in the Devonian (409–354 million years ago). Despite this, most scientists believe that
bryophytes evolved before vascular plants, and that the earliest bryophytes have not been found because they fossilize
poorly. This belief is supported by a variety of evidence, including morphological traits, ultrastructural features visible
under the electron microscope, and molecular information obtained from gene sequencing.
Bryophytes
Since bryophytes are land plants, they need to support themselves in air. However, because they lack lignified vascular
tissues, this support must be provided largely by the turgor pressure of their cells. Consequently, they cannot grow to be
very tall, and most bryophytes are small and rather inconspicuous. An additional important feature of their lifestyle is
their reproductive system. The male gametes , produced by reproductive structures called antheridia, are free-
swimming sperm cells that need water to transport them to the female gametes, which are enclosed within structures
called archegonia. Because of the need for water, bryophytes are especially common in wet habitats such as bogs,
streambanks, and in moist forests. However, they are not restricted to these habitats, and some mosses thrive in deserts,
above the treeline, and in the Arctic tundra.
Among the living bryophytes, liverworts are probably most closely related to the earliest land plants, since unlike
hornworts, mosses, and all vascular plants they do not possess stomata . Indeed, the fact that stomata first appeared in
hornworts and mosses is evidence that vascular plants evolved from one of these two groups. Vascular plants appear to
be more closely related to mosses than to hornworts, because some mosses possess food-conducting cells (leptoids) and
water-conducting cells (hydroids) that resemble the phloem and xylem of vascular plants.
Early Vascular Plants
The first detailed vascular plant fossils appear in rocks from middle Silurian, about 425 million years ago. The oldest of
these, including a plant called Aglaophyton, appear to have possessed conducting cells similar to the hydroids of
mosses. These ancient plants, which are sometimes called prototracheophytes, may have been an evolutionary link
between the bryophytes and the true tracheophytes. Early vascular plants possessed two features that made them
especially well adapted to life on land. First, their vascular tissues transported sugars, nutrients, and water far more
efficiently than the conducting cells of mosses. Second, they evolved the ability to synthesize lignin , which made the
cell walls of their vascular tissues rigid and supportive. Taken together, these features allowed them to grow much
larger than their bryophyte ancestors and considerably reduced their dependence on moist habitats.
There are three major groups of tracheophytes: seedless vascular plants, gymnosperms, and angiosperms. Since the first
appearance of tracheophytes in the Silurian, the fossil record shows three major evolutionary transitions, in each of
which a group of plants that were predominant before the transition is largely replaced by a different group that
becomes predominant afterward. The first such transition occurred in the late Devonian, approximately 375 million
years ago. Prior to this time the most common plants were simple, seedless vascular plants in various phyla, several of
which are now extinct. However, one phylum from this time, the Psilophyta, still has two living genera, including a
greenhouse weed called Psilotum.
From the late Devonian until the end of the Carboniferous period (290 million years ago) larger, more complex
seedless plants were predominant. The main phyla were the Lycophyta, the Sphenophyta, and the Pterophyta. All three
groups contain living relatives, including club mosses (Lycopodiaceae) in the Lycophyta, Equisetum (the only living
genus of sphenophytes), and ferns, which are pterophytes. Only the ferns, which have about 11,000 living species, are
common today, but in the Carboniferous these three phyla comprised a large fraction of the vegetation on the planet.
Many grew to the size of trees and dominated the tropical and subtropical swamps that covered much of the globe at
this time.
The second major transition was the decline of the lycophytes, sphenophytes, and pterophytes at the end of the
Carboniferous and their replacement by gymnosperms in the early Permian. Gymnosperms dominated the vegetation of
the land for the next 200 million years until they themselves began to decline and were replaced by angiosperms in the
middle of the Cretaceous. Although one group of gymnosperms (the conifers) is still abundant, the angiosperms have
been the most diverse and widespread group of plants on Earth for the last 100 million years.
Gymnosperms
The gymnosperms probably evolved from an extinct phylum of seedless vascular plants, the progymnosperms, that
appeared about 380 million years ago. The fossils of these plants, some of which were large trees, appear to form a link
between the trimerophytes (another extinct phylum of seedless vascular plants) and true gymnosperms.
Progymnosperms reproduced by means of spores like the former, but their vascular tissues were very similar to those of
living conifers. The oldest true gymnosperms, which produce seeds rather than spores, first appeared about 365 million
years ago. The evolution of seeds, with their hard, resilient coats, was almost certainly a key factor in the success of the
group. A second factor was the evolution of pollen grains to protect and transport the male gametes. As a consequence
of this, gymnosperms, unlike seedless vascular plants, were no longer dependent on water for successful fertilization
and could broadcast their male gametes on the wind.
Several early gymnosperm groups are now extinct, but there are four phyla with living representatives: the cycads, the
gnetophytes, the conifers, and one phylum (Ginkgophyta) that has only a single living species, the ginkgo tree (Ginkgo
biloba ). Of these, the conifers are by far the most abundant and diverse, and many species are of considerable
ecological and economic importance. Most conifers are well adapted to dry environments, particularly in their leaf
morphology , and some can withstand severe cold. These features may have enabled them to thrive in the Permian,
when Earth became much drier and colder than it had been in the Carboniferous.
Angiosperms
The angiosperms, or flowering plants, are all members of the phylum Anthophyta. There are at least 250,000 species,
making the group easily the most diverse of all plant phyla. They share a number of features that distinguish them from
other plant groups. The most obvious of these is the possession of flowers, highly modified shoots that carry the male
and female reproductive structures. They also carry out a process called double fertilization, in which two male
gametes (sperm nuclei) are released from the pollen tube into the ovule . One of these sperm nuclei fuses with an egg
cell in a similar way to gymnosperms. The second nucleus (which degenerates in most gymnosperms) fertilizes other
cells in the ovule called polar nuclei. Most commonly, two polar nuclei fuse with the sperm nucleus to form a triploid
endosperm nucleus. The tissue that forms from this fusion is called endosperm, which in most angiosperms provides
nutrients for the developing embryo.
A third feature that separates angiosperms from gymnosperms is that angiosperm embryos are protected by an ovary
wall, which develops into a fruit after fertilization has taken place. In contrast, gymnosperm embryos are held relatively
unprotected on the surfaces of ovule-bearing scales in the female cones.
Angiosperm Evolution
Angiosperms first appear in the fossil record about 130 million years ago, and by 90 million years ago they had become
the predominant group of plants on the planet. English naturalist Charles Darwin considered the sudden appearance of
angiosperms to be an "abominable mystery," and scientists have debated about the origin of the group for many years.
Comparative studies of living species suggest that angiosperms evolved from the gnetophytes, a group of gymnosperms
with three living genera of rather strange plants: Ephedra, Gnetum, and Welwitschia. Double fertilization has been
shown to occur in both Ephedra and Gnetum, and the reproductive structures (strobili) of all three genera are similar to
the flowering stalks of some angiosperms. Some gene sequencing studies also indicate that gnetophytes and
angiosperms are closely related to each other and to an extinct group of gymnosperms called the Bennettitales.
However, more recent molecular studies suggest that gnetophytes are more closely related to conifers than they are to
angiosperms.
In 1998, the discovery of an angiosperm-like fossil called Archaefructus, which apparently existed 145 million years
ago, also cast some doubt on the idea that angiosperms descended from gnetophytes or Bennettitales. Although a great
deal of information has been obtained since the time of Darwin, the origin of angiosperms is still something of a
mystery.
Early Angiosperms, Monocots, And Eudicots
The oldest known angiosperms were a diverse group of plants called magnoliids. Some of these were herbs with simple
flowers; others were woody plants with more complex flowers that were very similar to living magnolias. Magnoliids,
probably those with small, inconspicuous flowers, gave rise to the two main groups of angiosperms, monocots and
eudicots , although a few angiosperm families, including the water lilies, may have evolved earlier.
These plants possessed a number of adaptations that were probably crucial to their eventual success. Their vascular
tissues were particularly efficient, their embryos were enclosed in a protective seed coat, their leaves were resistant to
desiccation , and they were pollinated by insects, rather than by the wind. This last feature made pollen transfer much
more efficient and was almost certainly a key innovation in the diversification of the group, as coevolution of plants
and their pollinators, particularly bees, gave rise to increasing specialization of both flowers and insects.
The orchid family contains some of the most specialized insect-pollinated flowers of all and has more species (at least
24,000) than any other plant family. Other groups of angiosperms re-evolved the ability to be pollinated by wind. One
of these groups—the grasses—appeared about 50 million years ago, diversified rapidly, and became the dominant
plants over many regions of the planet. They still thrive and are crucial to human well-being. Approximately 54 percent
of the food eaten by people is provided by grain (seed) from cultivated varieties of just three grasses: rice, wheat, and
corn.
In the Beginning
Today we take for granted that we live among diverse communities of animals that feed on each other. Our ecosystems
are structured by feeding relationships like killer whales eating seals, which eat squid, which feed on krill. These and
other animals require oxygen to extract energy from their food. But that’s not how life on Earth used to be.
With an environment devoid of oxygen and high in methane, for much of its history Earth would not have been a
welcoming place for animals. The earliest life forms we know of were microscopic organisms (microbes) that left
signals of their presence in rocks about 3.7 billion years old. The signals consisted of a type of carbon molecule that is
produced by living things.
Evidence of microbes was also preserved in the hard structures (“stromatolites”) they made, which date to 3.5 billion
years ago. Stromatolites are created as sticky mats of microbes trap and bind sediments into layers. Minerals precipitate
inside the layers, creating durable structures even as the microbes die off. Scientists study today’s, rare living
stromatolite reefs to better understand Earth’s earliest life forms.
An Oxygen Atmosphere
When cyanobacteria evolved at least 2.4 billion years ago, they set the stage for a remarkable transformation. They
became Earth’s first photo-synthesizers, making food using water and the Sun’s energy, and releasing oxygen as a
result. This catalyzed a sudden, dramatic rise in oxygen, making the environment less hospitable for other microbes that
could not tolerate oxygen.
Evidence for this Great Oxidation Event is recorded in changes in seafloor rocks. When oxygen is around, iron reacts
chemically with it (it gets oxidized) and gets removed from the system. Rocks dating to before the event are striped
with bands of iron. Rocks dating to after the event do not have iron bands, showing that oxygen was now in the picture.
After the initial pulse of oxygen, it stabilized at lower levels where it would remain for a couple billion years more. In
fact, as cyanobacteria died and drifted down through the water, the decomposition of their bodies probably reduced
oxygen levels. So, the ocean was still not a suitable environment for most lifeforms that need ample oxygen.
Multicellular Life
However, other innovations were occurring. While they can process lots of chemicals, microbes did not have the
specialized cells that are needed for complex bodies. Animal bodies have various cells –skin, blood, bone – which
contain organelles, each doing a distinct job. Microbes are just single cells with no organelles and no nuclei to package
their DNA.
Something revolutionary happened as microbes began living inside other microbes, functioning as organelles for them.
Mitochondria, the organelles that process food into energy, evolved from these mutually beneficial relationships. Also,
for the first time, DNA became packaged in nuclei. The new complex cells (“eukaryotic cells”) boasted specialized
parts playing specialized roles that supported the whole cell.
Cells also began living together, probably because certain benefits could be obtained. Groups of cells might be able to
feed more efficiently or gain protection from simply being bigger. Living collectively, cells began to support the needs
of the group by each cell doing a specific job. Some cells were tasked with making junctions to hold the group together,
while other cells made digestive enzymes that could break down food.
The First Animals
These clusters of specialized, cooperating cells eventually became the first animals, which DNA evidence suggests
evolved around 800 million years ago. Sponges were among the earliest animals. While chemical compounds from
sponges are preserved in rocks as old as 700 million years, molecular evidence points to sponges developing even
earlier.
Oxygen levels in the ocean were still low compared to today, but sponges are able to tolerate conditions of low oxygen.
Although, like other animals, they require oxygen to metabolize, they don’t need much because they are not very
active. They feed while sitting still by extracting food particles from water that is pumped through their bodies by
specialized cells.
The simple body plan of a sponge consists of layers of cells around water-filled cavities, supported by hard skeletal
parts. The evolution of ever more complex and diverse body plans would eventually lead to distinct groups of animals.
The assembly instructions for an animal’s body plan are in its genes. Some genes act like orchestra conductors,
controlling the expression of many other genes at specific places and times to correctly assemble the components.
While they were not played out immediately, there is evidence that parts of instructions for complex bodies were
present even in the earliest animals.
Thanks to their hard skeletons, sponges became the first reef builders on Earth. Scientists are working to understand the
evolution of the thousands of sponge species living on Earth today.
Ediacaran Biota
By about 580 million years ago (the Ediacaran Period) there was a proliferation of other organisms, in addition to
sponges. These varied seafloor creatures - with bodies shaped like fronds, ribbons, and even quilts - lived alongside
sponges for 80 million years. Their fossil evidence can be found in sedimentary rocks around the world.
However, the body plans of most Ediacaran animals did not look like modern groups. Scientists are using comparative
developmental evidence, has examined whether any of the fossilized Ediacaran animals were related to modern
animals.
By the end of the Ediacaran, oxygen levels rose, approaching levels sufficient to sustain oxygen-based life. The early
sponges may actually have helped boost oxygen by eating bacteria, removing them from the decomposition process.
Tracks of an organism named Dickinsonia costata suggest that it may have been moved along the sea bottom,
presumably feasting on mats of microbes.
The End-Ediacaran Extinction
However, about 541 million years ago, most of the Ediacaran creatures disappeared, signaling a major environmental
change that Douglas Erwin and other scientists are still working to understand. Evolving animal body plans, feeding
relationships, and environmental engineering may have played a role.
Burrows found in the fossil record, dating to the end of the Ediacaran, reveal that worm-like animals had begun to
excavate the ocean bottom. These early environmental engineers disturbed and maybe aerated the sediment, disrupting
conditions for other Ediacaran animals. As environmental conditions deteriorated for some animals, they improved for
others, potentially catalyzing a change-over in species.
The Cambrian Explosion
The Cambrian Period (541-485 million years ago) witnessed a wild explosion of new life forms. Along with new
burrowing lifestyles came hard body parts like shells and spines. Hard body parts allowed animals to more drastically
engineer their environments, such as digging burrows. A shift also occurred towards more active animals, with defined
heads and tails for directional movement to chase prey. Active feeding by well-armored animals like trilobites may
have further disrupted the sea floor that the soft Ediacaran creatures had lived on.
Unique feeding styles partitioned the environment, making room for more diversification of life. In 1909 the
Smithsonian’s fourth Secretary, Charles Doolittle Walcott, discovered the Burgess Shale fossils that revealed the
unprecedented biodiversity of Cambrian life. While Waptia scoured the ocean bottom, priapulid worms burrowed into
the sediment, Wiwaxia attached to sponges, and Anomalocaris cruised above.
Many of these odd-looking organisms were evolutionary experiments, such as the 5-eyed Opabinia. However, some
groups, such as the trilobites, thrived and dominated Earth for hundreds of millions of years but eventually went
extinct. Stromatolite reef-building bacteria also declined, and reefs made by organisms called brachiopods arose as
conditions on Earth continued to change. Today’s dominant reef-builders, the hard corals, did not emerge until a couple
hundred million years later
However, despite all the changes that were to come, by the end of the Cambrian nearly all existing animal types, or
phyla, (mollusks, arthropods, annelids, etc.) were established, and food webs were emerging, forming the foundation
for the ecosystems on Earth today.