Evolution
Flowers of Malus sylvestris (crab apple)
Further information: Evolutionary history of plants#Flowers
Fossilized spores suggest that higher plants (embryophytes) have lived on the land for at least 475 million years.[3] Early land plants reproduced sexually with flagellated, swimming sperm, like the green algae from which they evolved. An adaptation to terrestrialization was the development of upright meiosporangia for dispersal by spores to new habitats. This feature is lacking in the descendants of their nearest algal relatives, the Charophycean green algae. A later terrestrial adaptation took place with retention of the delicate, avascular sexual stage, the gametophyte, within the tissues of the vascular sporophyte. This occurred by spore germination within sporangia rather than spore release, as in non-seed plants. A current example of how this might have happened can be seen in the precocious spore germination in Sellaginella, the spike-moss. The result for the ancestors of angiosperms was encloAsing them in a case, the seed. The first seed bearing plants, like the ginkgo, andconifers (such as pines and firs), did not produce flowers. The pollen grains (males) ofGinkgo and cycads produce a pair of flagellated, mobile sperm cells that "swim" down the developing pollen tube to the female and her eggs.
The apparently sudden appearance of relatively modern flowers in the fossil record initially posed such a problem for the theory of evolution that it was called an "abominable mystery" by Charles Darwin.[4] However, the fossil record has considerably grown since the time of Darwin, and recently discovered angiosperm fossils such as Archaefructus, along with further discoveries of fossil gymnosperms, suggest how angiosperm characteristics may have been acquired in a series of steps. Several groups of extinct gymnosperms, in particular seed ferns, have been proposed as the ancestors of flowering plants, but there is no continuous fossil evidence showing exactly how flowers evolved. Some older fossils, such as the upper Triassic Sanmiguelia, have been suggested. Based on current evidence, some propose that the ancestors of the angiosperms diverged from an unknown group of gymnosperms during the lateTriassic (245–202 million years ago). A close relationship between angiosperms and gnetophytes, proposed on the basis ofmorphological evidence, has more recently been disputed on the basis of molecular evidence that suggest gnetophytes are instead more closely related to other gymnosperms.[citation needed]
The evolution of seed plants and later angiosperms appears to be the result of two distinct rounds of whole genome duplication events.[5] These occurred in 319 million years ago and 192 million years ago respectively.
The earliest known macrofossil confidently identified as an angiosperm, Archaefructus liaoningensis, is dated to about 125 million years BP (the Cretaceous period),[6] while pollen considered to be of angiosperm origin takes the fossil record back to about 130 million years BP. However, one study has suggested that the early-middle Jurassic plant Schmeissneria, traditionally considered a type of ginkgo, may be the earliest known angiosperm, or at least a close relative.[7] In addition, circumstantial chemical evidence has been found for the existence of angiosperms as early as 250 million years ago. Oleanane, a secondary metabolite produced by many flowering plants, has been found in Permian deposits of that age together with fossils of gigantopterids.[8][9] Gigantopterids are a group of extinct seed plants that share many morphological traits with flowering plants, although they are not known to have been flowering plants themselves.
Recent DNA analysis based on molecular systematics [10][11] showed that Amborella trichopoda, found on the Pacific island of New Caledonia, belongs to a sister group of the other flowering plants, and morphological studies [12] suggest that it has features that may have been characteristic of the earliest flowering plants.
The orders Amborellales, Nymphaeales, and Austrobaileyales diverged as separate lineages from the remaining angiosperm clade at a very early stage in flowering plant evolution.[13]
The great angiosperm radiation, when a great diversity of angiosperms appears in the fossil record, occurred in the mid-Cretaceous(approximately 100 million years ago). However, a study in 2007 estimated that the division of the five most recent (the genusCeratophyllum, the family Chloranthaceae, the eudicots, the magnoliids, and the monocots) of the eight main groups occurred around 140 million years ago.[14] By the late Cretaceous, angiosperms appear to have dominated environments formerly occupied by ferns andcycadophytes, but large canopy-forming trees replaced conifers as the dominant trees only close to the end of the Cretaceous 65 millions years ago or even later, at the beginning of the Tertiary.[15] The radiation of herbaceous angiosperm occurred much later.[16]Yet, many fossil plants recognizable as belonging to modern families (including beech, oak, maple, and magnolia) appeared already at late Cretaceous.
Two bees on a flower head of Creeping Thistle, Cirsium arvense
It is generally assumed that the function of flowers, from the start, was to involve mobile animals in their reproduction processes. That is, pollen can be scattered even if the flower is not brightly coloredor oddly shaped in a way that attracts animals; however, by expending the energy required to create such traits, angiosperms can enlist the aid of animals and, thus, reproduce more efficiently.
Island genetics provides one proposed explanation for the sudden, fully developed appearance of flowering plants. Island genetics is believed to be a common source of speciation in general, especially when it comes to radical adaptations that seem to have required inferior transitional forms. Flowering plants may have evolved in an isolated setting like an island or island chain, where the plants bearing them were able to develop a highly specialized relationship with some specific animal (a wasp, for example). Such a relationship, with a hypothetical wasp carrying pollen from one plant to another much the way fig wasps do today, could result in the development of a high degree ofspecialization in both the plant(s) and their partners. Note that the wasp example is not incidental;bees, which, it is postulated, evolved specifically due to mutualistic plant relationships, are descended from wasps.
Animals are also involved in the distribution of seeds. Fruit, which is formed by the enlargement of flower parts, is frequently a seed-dispersal tool that attracts animals to eat or otherwise disturb it, incidentally scattering the seeds it contains (see frugivory). While many such mutualistic relationships remain too fragile to survive competition and to spread widely, flowering proved to be an unusually effective means of reproduction, spreading (whatever its origin) to become the dominant form of land plant life.
Flower ontogeny uses a combination of genes normally responsible for forming new shoots.[17] The most primitive flowers are thought to have had a variable number of flower parts, often separate from (but in contact with) each other. The flowers would have tended to grow in a spiral pattern, to be bisexual (in plants, this means both male and female parts on the same flower), and to be dominated by theovary (female part). As flowers grew more advanced, some variations developed parts fused together, with a much more specific number and design, and with either specific sexes per flower or plant, or at least "ovary-inferior".
Flower evolution continues to the present day; modern flowers have been so profoundly influenced by humans that some of them cannot be pollinated in nature. Many modern, domesticated flowers used to be simple weeds, which sprouted only when the ground was disturbed. Some of them tended to grow with human crops, perhaps already having symbiotic companion plant relationships with them, and the prettiest did not get plucked because of their beauty, developing a dependence upon and special adaptation to human affection.[18]
A few paleontologists have also come up with a theory that flowering plants, or angiosperms, might have evolved because of dinosaurs; in other words, they believe that dinosaurs "created" flowers. One of the theory's biggest proponents is Robert T. Bakker. He theorizes that herbivorous dinosaurs, with their eating habits, forced plants to find new ways to develop new adaptations, in order to avoid predation by herbivores.[citation needed]
Flowers of Malus sylvestris (crab apple)
Further information: Evolutionary history of plants#Flowers
Fossilized spores suggest that higher plants (embryophytes) have lived on the land for at least 475 million years.[3] Early land plants reproduced sexually with flagellated, swimming sperm, like the green algae from which they evolved. An adaptation to terrestrialization was the development of upright meiosporangia for dispersal by spores to new habitats. This feature is lacking in the descendants of their nearest algal relatives, the Charophycean green algae. A later terrestrial adaptation took place with retention of the delicate, avascular sexual stage, the gametophyte, within the tissues of the vascular sporophyte. This occurred by spore germination within sporangia rather than spore release, as in non-seed plants. A current example of how this might have happened can be seen in the precocious spore germination in Sellaginella, the spike-moss. The result for the ancestors of angiosperms was encloAsing them in a case, the seed. The first seed bearing plants, like the ginkgo, andconifers (such as pines and firs), did not produce flowers. The pollen grains (males) ofGinkgo and cycads produce a pair of flagellated, mobile sperm cells that "swim" down the developing pollen tube to the female and her eggs.
The apparently sudden appearance of relatively modern flowers in the fossil record initially posed such a problem for the theory of evolution that it was called an "abominable mystery" by Charles Darwin.[4] However, the fossil record has considerably grown since the time of Darwin, and recently discovered angiosperm fossils such as Archaefructus, along with further discoveries of fossil gymnosperms, suggest how angiosperm characteristics may have been acquired in a series of steps. Several groups of extinct gymnosperms, in particular seed ferns, have been proposed as the ancestors of flowering plants, but there is no continuous fossil evidence showing exactly how flowers evolved. Some older fossils, such as the upper Triassic Sanmiguelia, have been suggested. Based on current evidence, some propose that the ancestors of the angiosperms diverged from an unknown group of gymnosperms during the lateTriassic (245–202 million years ago). A close relationship between angiosperms and gnetophytes, proposed on the basis ofmorphological evidence, has more recently been disputed on the basis of molecular evidence that suggest gnetophytes are instead more closely related to other gymnosperms.[citation needed]
The evolution of seed plants and later angiosperms appears to be the result of two distinct rounds of whole genome duplication events.[5] These occurred in 319 million years ago and 192 million years ago respectively.
The earliest known macrofossil confidently identified as an angiosperm, Archaefructus liaoningensis, is dated to about 125 million years BP (the Cretaceous period),[6] while pollen considered to be of angiosperm origin takes the fossil record back to about 130 million years BP. However, one study has suggested that the early-middle Jurassic plant Schmeissneria, traditionally considered a type of ginkgo, may be the earliest known angiosperm, or at least a close relative.[7] In addition, circumstantial chemical evidence has been found for the existence of angiosperms as early as 250 million years ago. Oleanane, a secondary metabolite produced by many flowering plants, has been found in Permian deposits of that age together with fossils of gigantopterids.[8][9] Gigantopterids are a group of extinct seed plants that share many morphological traits with flowering plants, although they are not known to have been flowering plants themselves.
Recent DNA analysis based on molecular systematics [10][11] showed that Amborella trichopoda, found on the Pacific island of New Caledonia, belongs to a sister group of the other flowering plants, and morphological studies [12] suggest that it has features that may have been characteristic of the earliest flowering plants.
The orders Amborellales, Nymphaeales, and Austrobaileyales diverged as separate lineages from the remaining angiosperm clade at a very early stage in flowering plant evolution.[13]
The great angiosperm radiation, when a great diversity of angiosperms appears in the fossil record, occurred in the mid-Cretaceous(approximately 100 million years ago). However, a study in 2007 estimated that the division of the five most recent (the genusCeratophyllum, the family Chloranthaceae, the eudicots, the magnoliids, and the monocots) of the eight main groups occurred around 140 million years ago.[14] By the late Cretaceous, angiosperms appear to have dominated environments formerly occupied by ferns andcycadophytes, but large canopy-forming trees replaced conifers as the dominant trees only close to the end of the Cretaceous 65 millions years ago or even later, at the beginning of the Tertiary.[15] The radiation of herbaceous angiosperm occurred much later.[16]Yet, many fossil plants recognizable as belonging to modern families (including beech, oak, maple, and magnolia) appeared already at late Cretaceous.
Two bees on a flower head of Creeping Thistle, Cirsium arvense
It is generally assumed that the function of flowers, from the start, was to involve mobile animals in their reproduction processes. That is, pollen can be scattered even if the flower is not brightly coloredor oddly shaped in a way that attracts animals; however, by expending the energy required to create such traits, angiosperms can enlist the aid of animals and, thus, reproduce more efficiently.
Island genetics provides one proposed explanation for the sudden, fully developed appearance of flowering plants. Island genetics is believed to be a common source of speciation in general, especially when it comes to radical adaptations that seem to have required inferior transitional forms. Flowering plants may have evolved in an isolated setting like an island or island chain, where the plants bearing them were able to develop a highly specialized relationship with some specific animal (a wasp, for example). Such a relationship, with a hypothetical wasp carrying pollen from one plant to another much the way fig wasps do today, could result in the development of a high degree ofspecialization in both the plant(s) and their partners. Note that the wasp example is not incidental;bees, which, it is postulated, evolved specifically due to mutualistic plant relationships, are descended from wasps.
Animals are also involved in the distribution of seeds. Fruit, which is formed by the enlargement of flower parts, is frequently a seed-dispersal tool that attracts animals to eat or otherwise disturb it, incidentally scattering the seeds it contains (see frugivory). While many such mutualistic relationships remain too fragile to survive competition and to spread widely, flowering proved to be an unusually effective means of reproduction, spreading (whatever its origin) to become the dominant form of land plant life.
Flower ontogeny uses a combination of genes normally responsible for forming new shoots.[17] The most primitive flowers are thought to have had a variable number of flower parts, often separate from (but in contact with) each other. The flowers would have tended to grow in a spiral pattern, to be bisexual (in plants, this means both male and female parts on the same flower), and to be dominated by theovary (female part). As flowers grew more advanced, some variations developed parts fused together, with a much more specific number and design, and with either specific sexes per flower or plant, or at least "ovary-inferior".
Flower evolution continues to the present day; modern flowers have been so profoundly influenced by humans that some of them cannot be pollinated in nature. Many modern, domesticated flowers used to be simple weeds, which sprouted only when the ground was disturbed. Some of them tended to grow with human crops, perhaps already having symbiotic companion plant relationships with them, and the prettiest did not get plucked because of their beauty, developing a dependence upon and special adaptation to human affection.[18]
A few paleontologists have also come up with a theory that flowering plants, or angiosperms, might have evolved because of dinosaurs; in other words, they believe that dinosaurs "created" flowers. One of the theory's biggest proponents is Robert T. Bakker. He theorizes that herbivorous dinosaurs, with their eating habits, forced plants to find new ways to develop new adaptations, in order to avoid predation by herbivores.[citation needed]
