Flowering plant
| Flowering plants | |
|---|---|
 | |
| Diversity of angiosperms | |
Scientific classification | |
| Kingdom: | Plantae |
| Clade: | Tracheophytes |
| Clade: | Spermatophytes |
| Clade: | Angiosperms |
| Groups (APG IV)[1] | |
| |
| Synonyms | |
The flowering plants, also known as angiosperms, Angiospermae,[5][6] Magnoliophyta,[7] or, more colloquially, flowers, are the most diverse group of land plants, with 64 orders, 416 families, approximately 13,000 known genera and 300,000 known species.[8] Like gymnosperms, angiosperms are seed-producing plants. However, they are distinguished from gymnosperms by characteristics including flowers, endosperm within the seeds, and the production of fruits that contain the seeds. Etymologically, angiosperm means a plant that produces seeds within an enclosure; in other words, a fruiting plant. The term comes from the Greek words angeion ("case" or "casing") and sperma ("seed").
The ancestors of flowering plants diverged from gymnosperms in the Triassic Period, 245 to 202 million years ago (mya), and the first flowering plants are known from ~140 mya. They diversified extensively during the Early Cretaceous, became widespread by 120 mya, and replaced conifers as the dominant trees from 100 to 60 mya.
Contents
Description[edit]
Angiosperm derived characteristics[edit]
Angiosperms differ from other seed plants in several ways, described in the table below. These distinguishing characteristics taken together have made the angiosperms the most diverse and numerous land plants and the most commercially important group to humans.[a]
| Feature | Description |
|---|---|
| Flowering organs | Flowers, the reproductive organs of flowering plants, are the most remarkable feature distinguishing them from the other seed plants. Flowers provided angiosperms with the means to have a more species-specific breeding system, and hence a way to evolve more readily into different species without the risk of crossing back with related species. Faster speciation enabled the Angiosperms to adapt to a wider range of ecological niches. This has allowed flowering plants to largely dominate terrestrial ecosystems.[citation needed] |
| Stamens with two pairs of pollen sacs | Stamens are much lighter than the corresponding organs of gymnosperms and have contributed to the diversification of angiosperms through time with adaptations to specialized pollination syndromes, such as particular pollinators. Stamens have also become modified through time to prevent self-fertilization, which has permitted further diversification, allowing angiosperms eventually to fill more niches. |
| Reduced male parts, three cells | The male gametophyte in angiosperms is significantly reduced in size compared to those of gymnosperm seed plants.[9] The smaller size of the pollen reduces the amount of time between pollination — the pollen grain reaching the female plant — and fertilization. In gymnosperms, fertilization can occur up to a year after pollination, whereas in angiosperms, fertilization begins very soon after pollination.[10] The shorter amount of time between pollination and fertilization allows angiosperms to produce seeds earlier after pollination than gymnosperms, providing angiosperms a distinct evolutionary advantage. |
| Closed carpelenclosing the ovules (carpel or carpels and accessory parts may become the fruit) | The closed carpel of angiosperms also allows adaptations to specialized pollination syndromes and controls. This helps to prevent self-fertilization, thereby maintaining increased diversity. Once the ovary is fertilized, the carpel and some surrounding tissues develop into a fruit. This fruit often serves as an attractant to seed-dispersing animals. The resulting cooperative relationship presents another advantage to angiosperms in the process of dispersal. |
| Reduced female gametophyte, seven cells with eight nuclei | The reduced female gametophyte, like the reduced male gametophyte, may be an adaptation allowing for more rapid seed set, eventually leading to such flowering plant adaptations as annual herbaceous life-cycles, allowing the flowering plants to fill even more niches. |
| Endosperm | In general, endosperm formation begins after fertilization and before the first division of the zygote. Endosperm is a highly nutritive tissue that can provide food for the developing embryo, the cotyledons, and sometimes the seedlingwhen it first appears. |
Vascular anatomy[edit]
Angiosperm stems are made up of seven layers as shown on the right. The amount and complexity of tissue-formation in flowering plants exceeds that of gymnosperms. The vascular bundles of the stem are arranged such that the xylem and phloem form concentric rings.
In the dicotyledons, the bundles in the very young stem are arranged in an open ring, separating a central pith from an outer cortex. In each bundle, separating the xylem and phloem, is a layer of meristem or active formative tissue known as cambium. By the formation of a layer of cambium between the bundles (interfascicular cambium), a complete ring is formed, and a regular periodical increase in thickness results from the development of xylem on the inside and phloem on the outside. The soft phloem becomes crushed, but the hard wood persists and forms the bulk of the stem and branches of the woody perennial. Owing to differences in the character of the elements produced at the beginning and end of the season, the wood is marked out in transverse section into concentric rings, one for each season of growth, called annual rings.
Among the monocotyledons, the bundles are more numerous in the young stem and are scattered through the ground tissue. They contain no cambium and once formed the stem increases in diameter only in exceptional cases.
Reproductive anatomy[edit]
The characteristic feature of angiosperms is the flower. Flowers show remarkable variation in form and elaboration, and provide the most trustworthy external characteristics for establishing relationships among angiosperm species. The function of the flower is to ensure fertilization of the ovule and development of fruitcontaining seeds. The floral apparatus may arise terminally on a shoot or from the axil of a leaf (where the petiole attaches to the stem). Occasionally, as in violets, a flower arises singly in the axil of an ordinary foliage-leaf. More typically, the flower-bearing portion of the plant is sharply distinguished from the foliage-bearing or vegetative portion, and forms a more or less elaborate branch-system called an inflorescence.
There are two kinds of reproductive cells produced by flowers. Microspores, which will divide to become pollen grains, are the "male" cells and are borne in the stamens (or microsporophylls). The "female" cells called megaspores, which will divide to become the egg cell (megagametogenesis), are contained in the ovuleand enclosed in the carpel (or megasporophyll).
The flower may consist only of these parts, as in willow, where each flower comprises only a few stamens or two carpels. Usually, other structures are present and serve to protect the sporophylls and to form an envelope attractive to pollinators. The individual members of these surrounding structures are known as sepals and petals (or tepals in flowers such as Magnolia where sepals and petals are not distinguishable from each other). The outer series (calyx of sepals) is usually green and leaf-like, and functions to protect the rest of the flower, especially the bud. The inner series (corolla of petals) is, in general, white or brightly colored, and is more delicate in structure. It functions to attract insect or bird pollinators. Attraction is effected by color, scent, and nectar, which may be secreted in some part of the flower. The characteristics that attract pollinators account for the popularity of flowers and flowering plants among humans.
While the majority of flowers are perfect or hermaphrodite (having both pollen and ovule producing parts in the same flower structure), flowering plants have developed numerous morphological and physiological mechanisms to reduce or prevent self-fertilization. Heteromorphic flowers have short carpels and long stamens, or vice versa, so animal pollinators cannot easily transfer pollen to the pistil (receptive part of the carpel). Homomorphic flowers may employ a biochemical (physiological) mechanism called self-incompatibility to discriminate between self and non-self pollen grains. In other species, the male and female parts are morphologically separated, developing on different flowers.
Taxonomy[edit]
History of classification[edit]
The botanical term "Angiosperm", from the Ancient Greek ἀγγεῖον, angeíon (bottle, vessel) and σπέρμα, sperma (seed), was coined in the form Angiospermae by Paul Hermann in 1690, as the name of one of his primary divisions of the plant kingdom. This included flowering plants possessing seeds enclosed in capsules, distinguished from his Gymnospermae, or flowering plants with achenial or schizo-carpic fruits, the whole fruit or each of its pieces being here regarded as a seed and naked. The term and its antonym were maintained by Carl Linnaeus with the same sense, but with restricted application, in the names of the orders of his class Didynamia. Its use with any approach to its modern scope became possible only after 1827, when Robert Brown established the existence of truly naked ovules in the Cycadeae and Coniferae,[11] and applied to them the name Gymnosperms.[citation needed] From that time onward, as long as these Gymnosperms were, as was usual, reckoned as dicotyledonous flowering plants, the term Angiosperm was used antithetically by botanical writers, with varying scope, as a group-name for other dicotyledonous plants.
In 1851, Hofmeister discovered the changes occurring in the embryo-sac of flowering plants, and determined the correct relationships of these to the Cryptogamia. This fixed the position of Gymnosperms as a class distinct from Dicotyledons, and the term Angiosperm then gradually came to be accepted as the suitable designation for the whole of the flowering plants other than Gymnosperms, including the classes of Dicotyledons and Monocotyledons. This is the sense in which the term is used today.
In most taxonomies, the flowering plants are treated as a coherent group. The most popular descriptive name has been Angiospermae (Angiosperms), with Anthophyta ("flowering plants") a second choice. These names are not linked to any rank. The Wettstein system and the Engler system use the name Angiospermae, at the assigned rank of subdivision. The Reveal system treated flowering plants as subdivision Magnoliophytina(Frohne & U. Jensen ex Reveal, Phytologia 79: 70 1996), but later split it to Magnoliopsida, Liliopsida, and Rosopsida. The Takhtajan system and Cronquist system treat this group at the rank of division, leading to the name Magnoliophyta (from the family name Magnoliaceae). The Dahlgren system and Thorne system (1992) treat this group at the rank of class, leading to the name Magnoliopsida. The APG system of 1998, and the later 2003[12] and 2009[13] revisions, treat the flowering plants as a clade called angiosperms without a formal botanical name. However, a formal classification was published alongside the 2009 revision in which the flowering plants form the Subclass Magnoliidae.[14]
The internal classification of this group has undergone considerable revision. The Cronquist system, proposed by Arthur Cronquist in 1968 and published in its full form in 1981, is still widely used but is no longer believed to accurately reflect phylogeny. A consensus about how the flowering plants should be arranged has recently begun to emerge through the work of the Angiosperm Phylogeny Group (APG), which published an influential reclassification of the angiosperms in 1998. Updates incorporating more recent research were published as the APG II system in 2003,[12] the APG III system in 2009,[13][15] and the APG IV system in 2016.
Traditionally, the flowering plants are divided into two groups,
which in the Cronquist system are called Magnoliopsida (at the rank of class, formed from the family name Magnoliaceae) and Liliopsida (at the rank of class, formed from the family name Liliaceae). Other descriptive names allowed by Article 16 of the ICBN include Dicotyledones or Dicotyledoneae, and Monocotyledones or Monocotyledoneae, which have a long history of use. In English a member of either group may be called a dicotyledon (plural dicotyledons) and monocotyledon (plural monocotyledons), or abbreviated, as dicot (plural dicots) and monocot (plural monocots). These names derive from the observation that the dicots most often have two cotyledons, or embryonic leaves, within each seed. The monocots usually have only one, but the rule is not absolute either way. From a broad diagnostic point of view, the number of cotyledons is neither a particularly handy, nor a reliable character.
Recent studies, as by the APG, show that the monocots form a monophyletic group (clade) but that the dicots do not (they are paraphyletic). Nevertheless, the majority of dicot species do form a monophyletic group, called the eudicots or tricolpates. Of the remaining dicot species, most belong to a third major clade known as the magnoliids, containing about 9,000 species. The rest include a paraphyletic grouping of early branching taxa known collectively as the basal angiosperms, plus the families Ceratophyllaceae and Chloranthaceae.
Modern classification[edit]
There are eight groups of living angiosperms:
- Basal angiosperms (ANA: Amborella, Nymphaeales, Austrobaileyales)
- Amborella, a single species of shrub from New Caledonia;
- Nymphaeales, about 80 species,[16] water lilies and Hydatellaceae;
- Austrobaileyales, about 100 species[16] of woody plants from various parts of the world
- Core angiosperms (Mesangiospermae)[14]
- Chloranthales, 77 known species[17] of aromatic plants with toothed leaves;
- Magnoliids, about 9,000 species,[16] characterized by trimerous flowers, pollen with one pore, and usually branching-veined leaves—for example magnolias, bay laurel, and black pepper;
- Monocots, about 70,000 species,[16] characterized by trimerous flowers, a single cotyledon, pollen with one pore, and usually parallel-veined leaves—for example grasses, orchids, and palms;
- Ceratophyllum, about 6 species[16] of aquatic plants, perhaps most familiar as aquarium plants;
- Eudicots, about 175,000 species,[16] characterized by 4- or 5-merous flowers, pollen with three pores, and usually branching-veined leaves—for example sunflowers, petunia, buttercup, apples, and oaks.
The exact relationship between these eight groups is not yet clear, although there is agreement that the first three groups to diverge from the ancestral angiosperm were Amborellales, Nymphaeales, and Austrobaileyales.[18] The term basal angiosperms refers to these three groups. Among the remaining five groups (core angiosperms), the relationship between the three broadest of these groups (magnoliids, monocots, and eudicots) remains unclear. Zeng and colleagues (Fig. 1) describe four competing schemes.[19]Of these, eudicots and monocots are the largest and most diversified, with ~ 75% and 20% of angiosperm species, respectively. Some analyses make the magnoliids the first to diverge, others the monocots.[20] Ceratophyllum seems to group with the eudicots rather than with the monocots. The 2016 Angiosperm Phylogeny Group revision (APG IV) retained the overall higher order relationship described in APG III.[13]
|
|
|
| showDetailed Cladogram of the Angiosperm Phylogeny Group (APG) IV classification.[1] |
|---|
Evolutionary history[edit]
Fossilized spores suggest that land plants (embryophytes) have existed for at least 475 million years.[21] 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 Selaginella, the spike-moss. The result for the ancestors of angiosperms was enclosing them in a case, the seed. The first seed bearing plants, like the ginkgo, and conifers (such as pines and firs), did not produce flowers. The pollen grains (male gametophytes) of Ginkgo 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 nearly modern flowers in the fossil record initially posed such a problem for the theory of evolution that Charles Darwin called it an "abominable mystery".[22] 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 in the Triassic period (245–202 million years ago). Fossil angiosperm-like pollen from the Middle Triassic (247.2–242.0 Ma) suggests an older date for their origin.[23] A close relationship between angiosperms and gnetophytes, proposed on the basis of morphological evidence, has more recently been disputed on the basis of molecular evidence that suggest gnetophytes are instead more closely related to other gymnosperms.[24][25] The fossil plant species Nanjinganthus dendrostyla from Early Jurassic China seems to share many exclusively angiosperm features, such as a thickened receptacle with ovules, and thus might represent a crown-group or a stem-group angiosperm.[26][27] However, the interpretation of the structures in this fossils are highly contested.[28][29]
The evolution of seed plants and later angiosperms appears to be the result of two distinct rounds of whole genome duplication events.[30]These occurred at 319 million years ago and 192 million years ago. Another possible whole genome duplication event at 160 million years ago perhaps created the ancestral line that led to all modern flowering plants.[31] That event was studied by sequencing the genome of an ancient flowering plant, Amborella trichopoda,[32] and directly addresses Darwin's "abominable mystery".
The earliest known macrofossil confidently identified as an angiosperm, Archaefructus liaoningensis, is dated to about 125 million years BP(the Cretaceous period),[33] whereas pollen considered to be of angiosperm origin takes the fossil record back to about 130 million years BP.[34] 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.[35] In 2018, scientists reported that the earliest flowers began about 180 million years ago, 50 million years earlier than thought earlier.[36] Nonetheless, 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.[37][38] 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.[citation needed]
In 2013 flowers encased in amber were found and dated 100 million years before present. The amber had frozen the act of sexual reproduction in the process of taking place. Microscopic images showed tubes growing out of pollen and penetrating the flower's stigma. The pollen was sticky, suggesting it was carried by insects.[39]
Recent DNA analysis based on molecular systematics[40][41] 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[42] 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.[43]
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 genus Ceratophyllum, the family Chloranthaceae, the eudicots, the magnoliids, and the monocots) of the eight main groups occurred around 140 million years ago.[44] By the late Cretaceous, angiosperms appear to have dominated environments formerly occupied by ferns and cycadophytes, but large canopy-forming trees replaced conifers as the dominant trees only close to the end of the Cretaceous 66 million years ago or even later, at the beginning of the Tertiary.[45]The radiation of herbaceous angiosperms occurred much later.[46] Yet, many fossil plants recognizable as belonging to modern families (including beech, oak, maple, and magnolia) had already appeared by the late Cretaceous.
It has been proposed that the swift rise of angiosperms to dominance was facilitated by a reduction in their genome size. During the early Cretaceous period, only angiosperms underwent rapid genome downsizing, while genome sizes of ferns and gymnosperms remained unchanged. Smaller genomes—and smaller nuclei—allow for faster rates of cell division and smaller cells. Thus, species with smaller genomes can pack more, smaller cells—in particular veins and stomata—into a given leaf volume. Genome downsizing therefore facilitated higher rates of leaf gas exchange (transpiration and photosynthesis) and faster rates of growth. This would have countered some of the negative physiological effects of genome duplications, facilitated increased uptake of carbon dioxide despite concurrent declines in atmospheric CO2 concentrations, and allowed the flowering plants to outcompete other land plants.[47]
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 colored or 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 of specialization 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.[48]
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). Although 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.[49] The most primitive flowers probably had a variable number of flower parts, often separate from (but in contact with) each other. The flowers 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 the ovary (female part). As flowers evolved, 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 flower species were formerly 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.[50]
A few paleontologists have also proposed that flowering plants, or angiosperms, might have evolved due to interactions with dinosaurs. One of the idea's strongest proponents is Robert T. Bakker. He proposes that herbivorous dinosaurs, with their eating habits, provided a selective pressure on plants, for which adaptations either succeeded in deterring or coping with predation by herbivores.[51]
In August 2017, scientists presented a detailed description and 3D model image of what the first flower possibly looked like, and presented the hypothesis that it may have lived about 140 million years ago.[52][53]
A Bayesian analysis of 52 angiosperm taxa suggested that the crown group of angiosperms evolved between 178 million years ago and 198 million years ago.[54]
Diversity[edit]
The number of species of flowering plants is estimated to be in the range of 250,000 to 400,000.[55][56][57] This compares to around 12,000 species of moss[58] or 11,000 species of pteridophytes,[59] showing that the flowering plants are much more diverse. The number of familiesin APG (1998) was 462. In APG II[12] (2003) it is not settled; at maximum it is 457, but within this number there are 55 optional segregates, so that the minimum number of families in this system is 402. In APG III (2009) there are 415 families.[13][60]
The diversity of flowering plants is not evenly distributed. Nearly all species belong to the eudicot (75%), monocot (23%), and magnoliid (2%) clades. The remaining 5 clades contain a little over 250 species in total; i.e. less than 0.1% of flowering plant diversity, divided among 9 families. The 43 most-diverse of 443 families of flowering plants by species,[61] in their APG circumscriptions, are
- Asteraceae or Compositae (daisy family): 22,750 species;
- Orchidaceae (orchid family): 21,950;
- Fabaceae or Leguminosae (bean family): 19,400;
- Rubiaceae (madder family): 13,150;[62]
- Poaceae or Gramineae (grass family): 10,035;
- Lamiaceae or Labiatae (mint family): 7,175;
- Euphorbiaceae (spurge family): 5,735;
- Melastomataceae or Melastomaceae (melastome family): 5,005;
- Myrtaceae (myrtle family): 4,625;
- Apocynaceae (dogbane family): 4,555;
- Cyperaceae (sedge family): 4,350;
- Malvaceae (mallow family): 4,225;
- Araceae (arum family): 4,025;
- Ericaceae (heath family): 3,995;
- Gesneriaceae (gesneriad family): 3,870;
- Apiaceae or Umbelliferae (parsley family): 3,780;
- Brassicaceae or Cruciferae (cabbage family): 3,710:
- Piperaceae (pepper family): 3,600;
- Bromeliaceae (bromeliad family): 3,540;
- Acanthaceae (acanthus family): 3,500;
- Rosaceae (rose family): 2,830;
- Boraginaceae (borage family): 2,740;
- Urticaceae (nettle family): 2,625;
- Ranunculaceae (buttercup family): 2,525;
- Lauraceae (laurel family): 2,500;
- Solanaceae (nightshade family): 2,460;
- Campanulaceae (bellflower family): 2,380;
- Arecaceae (palm family): 2,361;
- Annonaceae (custard apple family): 2,220;
- Caryophyllaceae (pink family): 2,200;
- Orobanchaceae (broomrape family): 2,060;
- Amaranthaceae (amaranth family): 2,050;
- Iridaceae (iris family): 2,025;
- Aizoaceae or Ficoidaceae (ice plant family): 2,020;
- Rutaceae (rue family): 1,815;
- Phyllanthaceae (phyllanthus family): 1,745;
- Scrophulariaceae (figwort family): 1,700;
- Gentianaceae (gentian family): 1,650;
- Convolvulaceae (bindweed family): 1,600;
- Proteaceae (protea family): 1,600;
- Sapindaceae (soapberry family): 1,580;
- Cactaceae (cactus family): 1,500;
- Araliaceae (Aralia or ivy family): 1,450.
Of these, the Orchidaceae, Poaceae, Cyperaceae, Araceae, Bromeliaceae, Arecaceae, and Iridaceae are monocot families; Piperaceae, Lauraceae, and Annonaceae are magnoliid dicots; the rest of the families are eudicots.
Reproduction[edit]
Fertilization and embryogenesis[edit]
Double fertilization refers to a process in which two sperm cells fertilize cells in the ovule. This process begins when a pollen grain adheres to the stigma of the pistil (female reproductive structure), germinates, and grows a long pollen tube. While this pollen tube is growing, a haploid generative cell travels down the tube behind the tube nucleus. The generative cell divides by mitosis to produce two haploid (n) sperm cells. As the pollen tube grows, it makes its way from the stigma, down the style and into the ovary. Here the pollen tube reaches the micropyle of the ovule and digests its way into one of the synergids, releasing its contents (which include the sperm cells). The synergid that the cells were released into degenerates and one sperm makes its way to fertilize the egg cell, producing a diploid (2n) zygote. The second sperm cell fuses with both central cell nuclei, producing a triploid (3n) cell. As the zygote develops into an embryo, the triploid cell develops into the endosperm, which serves as the embryo's food supply. The ovary will now develop into a fruit and the ovule will develop into a seed.
Fruit and seed[edit]
As the development of embryo and endosperm proceeds within the embryo sac, the sac wall enlarges and combines with the nucellus (which is likewise enlarging) and the integument to form the seed coat. The ovary wall develops to form the fruit or pericarp, whose form is closely associated with type of seed dispersal system.[63]
Frequently, the influence of fertilization is felt beyond the ovary, and other parts of the flower take part in the formation of the fruit, e.g., the floral receptacle in the apple, strawberry, and others.[citation needed]
The character of the seed coat bears a definite relation to that of the fruit. They protect the embryo and aid in dissemination; they may also directly promote germination. Among plants with indehiscent fruits, in general, the fruit provides protection for the embryo and secures dissemination. In this case, the seed coat is only slightly developed. If the fruit is dehiscent and the seed is exposed, in general, the seed-coat is well developed, and must discharge the functions otherwise executed by the fruit.[citation needed]
Meiosis[edit]
Flowering plants generate gametes using a specialized cell division called meiosis. Meiosis takes place in the ovule (a structure within the ovary that is located within the pistil at the center of the flower) (see diagram labeled "Angiosperm lifecycle"). A diploid cell (megaspore mother cell) in the ovule undergoes meiosis (involving two successive cell divisions) to produce four cells (megaspores) with haploid nuclei.[64] One of these four cells (megaspore) then undergoes three successive mitotic divisions to produce an immature embryo sac (megagametophyte) with eight haploid nuclei. Next, these nuclei are segregated into separate cells by cytokinesis to producing 3 antipodal cells, 2 synergid cells and an egg cell. Two polar nuclei are left in the central cell of the embryo sac.[citation needed]
Pollen is also produced by meiosis in the male anther (microsporangium). During meiosis, a diploid microspore mother cell undergoes two successive meiotic divisions to produce 4 haploid cells (microspores or male gametes). Each of these microspores, after further mitoses, becomes a pollen grain (microgametophyte) containing two haploid generative (sperm) cells and a tube nucleus. When a pollen grain makes contact with the female stigma, the pollen grain forms a pollen tube that grows down the style into the ovary. In the act of fertilization, a male sperm nucleus fuses with the female egg nucleus to form a diploid zygote that can then develop into an embryo within the newly forming seed. Upon germination of the seed, a new plant can grow and mature.[citation needed]
The adaptive function of meiosis is currently a matter of debate. A key event during meiosis in a diploid cell is the pairing of homologous chromosomes and homologous recombination (the exchange of genetic information) between homologous chromosomes. This process promotes the production of increased genetic diversity among progeny and the recombinational repair of damages in the DNA to be passed on to progeny. To explain the adaptive function of meiosis in flowering plants, some authors emphasize diversity[65]and others emphasize DNA repair.[66]
Apomixis[edit]
Apomixis (reproduction via asexually formed seeds) is found naturally in about 2.2% of angiosperm genera.[67] One type of apomixis, gametophytic apomixis found in a dandelion species[68] involves formation of an unreduced embryo sac due to incomplete meiosis (apomeiosis) and development of an embryo from the unreduced egg inside the embryo sac, without fertilization (parthenogenesis).[citation needed]
Uses[edit]
Agriculture is almost entirely dependent on angiosperms, which provide virtually all plant-based food, and also provide a significant amount of livestock feed. Of all the families of plants, the Poaceae, or grass family (providing grains), is by far the most important, providing the bulk of all feedstocks (rice, maize, wheat, barley, rye, oats, pearl millet, sugar cane, sorghum). The Fabaceae, or legume family, comes in second place. Also of high importance are the Solanaceae, or nightshade family (potatoes, tomatoes, and peppers, among others); the Cucurbitaceae, or gourd family (including pumpkins and melons); the Brassicaceae, or mustard plant family (including rapeseed and the innumerable varieties of the cabbage species Brassica oleracea); and the Apiaceae, or parsley family. Many of our fruits come from the Rutaceae, or rue family (including oranges, lemons, grapefruits, etc.), and the Rosaceae, or rose family (including apples, pears, cherries, apricots, plums, etc.).[citation needed]
In some parts of the world, certain single species assume paramount importance because of their variety of uses, for example the coconut (Cocos nucifera) on Pacific atolls, and the olive (Olea europaea) in the Mediterranean region.[69]
Flowering plants also provide economic resources in the form of wood, paper, fiber (cotton, flax, and hemp, among others), medicines (digitalis, camphor), decorative and landscaping plants, and many other uses. The main area in which they are surpassed by other plants—namely, coniferous trees (Pinales), which are non-flowering (gymnosperms)— is timber and paper production
Also called Angiospermae. The ancestors of flowering plants diverged from gymnosperms around 245–202 million years ago, and the first flowering plants known to exist are from 140 million years ago. They diversified enormously during the Lower Cretaceous and became widespread around 100 million years ago, but replaced conifers as the dominant trees only around 60-100 million years ago.
The Division Magnoliophyta in the Kingdom Plantae comprises those species of plants that were formerly classified as angiosperms and are known widely as the flowering plants. You have already studied flowers (Chapter 4), so now understand that the Division Magnoliophyta comprises all those species of plants that have flowers. For perspective: all of the plants we have read about in Section II of the Botany Study Guide up to this point do not have flowers, but certainly do have reproductive structures. We also know that flowers are the reproductive structures of the plants that bear them, and that reproductive structures are not limited to flowering plants. Thus, "flowers" are structures that distinguish plants in the Division Magnoliophyta from plants in the other divisions of the Kingdom Plantae. Observe a flower, and you know you are examining an angiosperm. However, not all angiosperms have obvious, showy flowers. You will need to consider that the structure of a flower is quite variable across all the many species of angiosperms (about a quarter million have been identified) and a few flowering plants actually seldom flower. However, angiosperm botanists put great stock in the structure of flowers as a way of classifying plants—more than any other part of a plant, the flower provides the basis for placement of a species in subtaxa (classes, orders, and families) of the Division Magnoliophyta.
The Division Magnoliophyta is split into two large classes: the Magnoliopsida and the Liliopsida.
- Read Flowering plants (The following links are included)
- Reread cotyledon article
- Read dicotyledons (Magnoliopsida)
- Read monocotyledons (Liliopsida)
- Read How to distinguish monocots and dicots and follow any links that you are uncertain about
There are over 300 families and 250,000 species of flowering plants. The remainder of this chapter will be concerned with the dicotyledonous angiosperms (Class Magnoliopsida), leaving the monocotyledonous angiosperms to be covered in the next chapter.
Flower evolution[edit]
Conifers are pollinated by wind, meaning they must produce a large amount of pollen grains for only a few to arrive at the female megaspore, resulting in fertilization. An advancement that allowed for higher rates of fertilization per energy expended in making pollen would surely result in that new plant type being prolific and even dominant. However, a single mutation would never result in a flower adequately adapted to spread pollen using animals. Flowers are the result of a special kind of evolution called co-evolution.
If plants reproduced and thrived with wind pollination, why did flowers evolve?
Suppose one wind pollinated plant began to have its' pollen eaten by an animal, for example a beetle. Then suppose the beetle spreads the pollen from the male cones to the female cones while looking for pollen because it can't tell the difference between the male and female cones; it searches both, spreading pollen from the male to the female in doing so. This plant has an advantage over all wind pollinated plants, because of a higher rate of fertilization. Any mutation that leads to pollen being spread by an animal soon becomes prevalent within a species, because the plant pollinated by an animal is more efficient. The plant and the animal rely on each other, one for food, the other for reproduction. Any mutation in the animal that helps pollinate the plant will become prolific through natural selection, just as mutations that favor feeding the insect or making the insect come to the flower will be dominant. This back and forth evolution results in such seemingly improbable structures as flowers that look like female moths that are pollinated by amorous male moths, or flowers and bird beaks that complement the feeding of the bird and the pollination of the flowers.
Class Magnoliopsida (dicots)[edit]
Members of the Class Magnoliopsida are defined partly on the basis of the seed or seedling having two cotyledons, most obvious at germination. But the differences between dicots and monocots are many, and we will be able to recognize most flowering plants that we encounter as belonging to one or the other class without having to dissect the seed or observe the seedling.
Angiosperms, also called flowering plants, have seeds that are enclosed within an ovary (usually a fruit), while gymnosperms have no flowers or fruits, and have unenclosed or “naked” seeds on the surface of scales or leaves. Gymnosperm seeds are often configured as cones. The characteristics that differentiate angiosperms from gymnosperms include flowers, fruits, and endosperm in the seeds.
Comparison chart
 | Angiosperms | Gymnosperms |
|---|---|---|
| Definition | Seed-producing flowering plants whose seeds are enclosed within an ovary. | Seed-producing non-flowering plants whose seeds are unenclosed or “naked.” |
| Seeds | Enclosed inside an ovary, usually in a fruit. | Bare, not enclosed; found on scales, leaves or as cones. |
| Life Cycle | Seasonal (die during autumn/fall). | Evergreen |
| Tissue | Triploid (endosperm produced during triple fusion) | Haploid (endosperm produced before fertilization) |
| Reproductive system | Present in flowers; can be unisexual or bisexual | Cones; unisexual |
| Leaves | Flat | scalelike , needle-like |
| Cotyledons | Present; single (monocots) or in a pair (dicots) | Absent |
| Wood | Hardwood | Softwood |
| Periniality | Non-perinnial | Perinnial |
| Kingdom | Plantae | Plantae |
| Domain | Eukarya | Eukarya |
| Reproduction | Mostly rely on animals. | Mostly rely on wind. |
| Uses | Medications , food, clothing, etc... | Paper, Lumber , etc... |
Diversity
Hundreds of millions of years ago, gymnosperms were the only kind of plant life on Earth. Between 250 and 200 million years ago, angiosperms started to evolve. Now, angiosperms are more widely distributed and populous, and can be considered the dominant plant life on the planet. Angiosperms comprise a far more diverse range of plants, with a range of 250,000 to 400,000 species. They inhabit every kind of land and aquatic environment except the most extreme habitats. Angiosperms may be dicots or monocots.
Examples of Angiosperms and Gymnosperms
Examples of angiosperms are monocots like lilies, orchids, agaves (known for agave nectar) and grasses; and dicots like roses, peas, sunflowers, oaks and maples.
Gymnosperm examples include non-flowering evergreen trees such as pine, spruce and fir.
Gymnosperm species number only in the thousands, with a little more than 1,000 extant species. They are found in desert to semi-desert habitats.
Anatomical Differences
Since gymnosperms and angiosperms are both vascular plants, they have a sporophyte-dominant life-cycle.
Tissue formation in angiosperms exceeds the amount and complexity found in gymnosperms. Angiosperms have a triploid vascular tissue, flat leaves in numerous shapes and hardwood stems. Because of the innumerable varieties of the fruit and/or flower-bearing plants, they have variegated colors and shapes of leaves, flowers and fruits.
Gymnosperms are haploid, have spiky, needle-like leaves and are softwood. Gymnosperms are "simpler" anatomically because they do not bear flowers or fruit, and although of different species, are usually only tall evergreens with brown cones.
More details about the anatomical differences between angiosperms and gymnosperms are explained in the following video:
Reproduction
Reproduction in angiosperms can be unisexual or bisexual. The gametes are spread by wind and by insect and animal pollinators attracted by their flowers. Flowers often have both female and male gametes inside them, and after fertilization, the ovules develop into a fruit.
The gametes of gymnosperms are found in cones. Fertilization is described as single; the pollen grains fall and germinate directly on the ovules. Pollen spores are spread by wind alone.
Uses
Angiosperms provide virtually all plant-based food, as well as most livestock feed. Grains, fruit, legumes, nightshades (including potatoes and tomatoes), gourds, and cabbages are all angiosperms. Other angiosperms like cotton and flax provide paper and textiles. The hardwood of angiosperms is used to make hardwood floors.
Gymnosperms from the conifer group like pine, spruce, and fir are commonly used for lumber. Other gymnosperms are processed into other products like soap, varnish, and perfumes.