Evolution of Plants
Photo by: Andrey Armyagov
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 particular 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.
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.
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.
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
The ginkgo tree ( Ginkgo biloba ) is the only living species of the early gymnosperm phylum Ginkgophyta.
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.
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.
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Evolution, Evidence for Excretory Systems
The eudicots, Eudicotidae or eudicotyledons are a clade of flowering plants that had been called tricolpates or non-magnoliid dicots by previous authors. The botanical terms were introduced in 1991 by evolutionary botanist James A. Doyle and paleobotanist Carol L. Hotton to emphasize the later evolutionary divergence of tricolpate dicots from earlier, less specialized, dicots. The close relationships among flowering plants with tricolpate pollen grains was initially seen in morphological studies of shared derived characters. These plants have a distinct trait in their pollen grains of exhibiting three colpi or grooves paralleling the polar axis. Later molecular evidence confirmed the genetic basis for the evolutionary relationships among flowering plants with tricolpate pollen grains and dicotyledonous traits. The term means "true dicotyledons", as it contains the majority of plants that have been considered dicots and have characteristics of the dicots. The term "eudicots" has subsequently been widely adopted in botany to refer to one of the two largest clades of angiosperms (constituting over 70% of the angiosperm species), monocots being the other. The remaining angiosperms include magnoliids and what are sometimes referred to as basal angiosperms or paleodicots, but these terms have not been widely or consistently adopted, as they do not refer to a monophyletic group.
The other name for the eudicots is tricolpates, a name which refers to the grooved structure of the pollen. Members of the group have tricolpate pollen, or forms derived from it. These pollens have three or more pores set in furrows called colpi. In contrast, most of the other seed plants (that is the gymnosperms, the monocots and the paleodicots) produce monosulcate pollen, with a single pore set in a differently oriented groove called the sulcus. The name "tricolpates" is preferred by some botanists to avoid confusion with the dicots, a nonmonophyletic group.
Numerous familiar plants are eudicots, including many common food plants, trees, and ornamentals. Some common and familiar eudicots include members of the sunflower family such as the common dandelion, the forget-me-not, cabbage and other members of its family, apple, buttercup, maple, and macadamia. Most leafy trees of midlatitudes also belong to eudicots, with notable exceptions being magnolias and tulip trees which belong to magnoliids, and Ginkgo biloba, which is not an angiosperm.
The name "eudicots" (plural) is used in the APG system, of 1998, and APG II system, of 2003, for classification of angiosperms. It is applied to a clade, a monophyletic group, which includes most of the (former) dicots.
The eudicots can be divided into two groups: the basal eudicots and the core eudicots. Basal eudicot is an informal name for a paraphyletic group. The core eudicots are a monophyletic group. A 2010 study suggested the core eudicots can be divided into two clades, Gunnerales and a clade called "Pentapetalae", comprising all the remaining core eudicots.
The Pentapetalae can be then divided into three clades:
This division of the eudicots is shown in the following cladogram:
The following is a more detailed breakdown according to APG IV, showing within each clade and orders:
- ^Endress, Peter K. (2002). "Morphology and Angiosperm Systematics in the Molecular Era". Botanical Review. Structural Botany in Systematics: A Symposium in Memory of William C. Dickison. New York: New York Botanical Garden Press. 68 (4): 545–570. doi:10.1663/0006-8101(2002)068[0545:maasit]2.0.co;2. ISSN 0006-8101. JSTOR 4354438.
- ^(Judd & Olmstead 2004).
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Stevens, P.F. (2001–2014). "Trees". Angiosperm Phylogeny Website. Retrieved 2014-11-17.
Stevens, P.F. (2001–2016). "Eudicots". Angiosperm Phylogeny Website. Retrieved 2014-11-17.
- ^Angiosperm Phylogeny Group (2016). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV"(PDF). Botanical Journal of the Linnean Society. 181 (1): 1–20. doi:10.1111/boj.12385. Retrieved 2016-04-10.
- Doyle, J. A. & Hotton, C. L. Diversification of early angiosperm pollen in a cladistic context. pp. 169–195 in Pollen and Spores. Patterns of Diversification (eds Blackmore, S. & Barnes, S. H.) (Clarendon, Oxford, 1991).
- Walter S. Judd and Richard G. Olmstead (2004). "A survey of tricolpate (eudicot) phylogenetic relationships". American Journal of Botany. 91 (10): 1627–1644. doi:10.3732/ajb.91.10.1627. PMID 21652313. (full text )
- Eudicots in Stevens, P. F. (2001 onwards). Angiosperm Phylogeny Website. Version 7, May 2006.
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