Flowering plant

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Flowering plants
Temporal range: Early Cretaceouspresent, 130–0  Ma
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Possible Early Jurassic record
Flower poster 2.jpg
Diversity of Angiosperms
Scientific classification Red Pencil Icon.png
Kingdom: Plantae
Clade: Spermatophytes
Clade:Angiosperms
Groups (APG IV) [1]

Nanjinganthus ?

Basal angiosperms

Core angiosperms

Synonyms

The flowering plants, also known as angiosperms, Angiospermae [5] [6] or Magnoliophyta, [7] 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").

Embryophyte Subkingdom of plants

The Embryophyta, or land plants, are the most familiar group of green plants that form vegetation on earth. Embryophyta is a clade within the Phragmoplastophyta, a larger clade that also includes several green algae groups, and within this large clade the embryophytes are sister to the Zygnematophyceae/Mesotaeniaceae and consist of the bryophytes plus the polysporangiophytes. Living embryophytes therefore include hornworts, liverworts, mosses, ferns, lycophytes, gymnosperms and flowering plants. The Embryophyta are informally called land plants because they live primarily in terrestrial habitats, while the related green algae are primarily aquatic. Embryophytes are complex multicellular eukaryotes with specialized reproductive organs. The name derives from their innovative characteristic of nurturing the young embryo sporophyte during the early stages of its multicellular development within the tissues of the parent gametophyte. With very few exceptions, embryophytes obtain their energy by photosynthesis, that is by using the energy of sunlight to synthesize their food from carbon dioxide and water.

Family is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy; it is classified between order and genus. A family may be divided into subfamilies, which are intermediate ranks between the ranks of family and genus. The official family names are Latin in origin; however, popular names are often used: for example, walnut trees and hickory trees belong to the family Juglandaceae, but that family is commonly referred to as being the "walnut family".

In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. Other ways of defining species include their karyotype, DNA sequence, morphology, behaviour or ecological niche. In addition, paleontologists use the concept of the chronospecies since fossil reproduction cannot be examined. While these definitions may seem adequate, when looked at more closely they represent problematic species concepts. For example, the boundaries between closely related species become unclear with hybridisation, in a species complex of hundreds of similar microspecies, and in a ring species. Also, among organisms that reproduce only asexually, the concept of a reproductive species breaks down, and each clone is potentially a microspecies.

Contents

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 160 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.

The Triassic is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.9 million years ago (Mya), to the beginning of the Jurassic Period 201.3 Mya. The Triassic is the first and shortest period of the Mesozoic Era. Both the start and end of the period are marked by major extinction events.

The abbreviation myr, "million years", is a unit of a quantity of 1,000,000 (i.e. 1×106) years, or 31.536 teraseconds.

The Early Cretaceous or the Lower Cretaceous, is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 146 Ma to 100 Ma.

Description

Angiosperm derived characteristics

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. [lower-alpha 1]

Spermatophyte division of plants

The spermatophytes, also known as phanerogams or phaenogams, comprise those plants that produce seeds, hence the alternative name seed plants. They are a subset of the embryophytes or land plants. The term phanerogams or phanerogamae is derived from the Greek φανερός, phanerós meaning "visible", in contrast to the cryptogamae from Greek κρυπτός kryptós = "hidden" together with the suffix γαμέω, gameein, "to marry". These terms distinguished those plants with hidden sexual organs (cryptogamae) from those with visible sexual organs (phanerogamae).

Distinctive features of angiosperms
FeatureDescription
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 sacsStamens 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 carpel enclosing 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 nucleiThe 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 seedling when it first appears.

Vascular anatomy

Cross-section of a stem of the angiosperm flax:
1. pith, 2. protoxylem, 3. xylem, 4. phloem, 5. sclerenchyma (bast fibre), 6. cortex, 7. epidermis Stem-histology-cross-section-tag.svg
Cross-section of a stem of the angiosperm flax:
1. pith, 2. protoxylem, 3. xylem, 4. phloem, 5. sclerenchyma (bast fibre), 6. cortex, 7. epidermis

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.

Complexity characterises the behaviour of a system or model whose components interact in multiple ways and follow local rules, meaning there is no reasonable higher instruction to define the various possible interactions.

Vascular bundle part of a plant

A vascular bundle is a part of the transport system in vascular plants. The transport itself happens in vascular tissue, which exists in two forms: xylem and phloem. Both these tissues are present in a vascular bundle, which in addition will include supporting and protective tissues.

Xylem One of the two types of transport tissue in vascular plants. xylems transport water from roots to shoots and leaves, but also some nutrients

Xylem is one of the two types of transport tissue in vascular plants, phloem being the other. The basic function of xylem is to transport water from roots to stems and leaves, but it also transports nutrients. The word "xylem" is derived from the Greek word ξύλον (xylon), meaning "wood"; the best-known xylem tissue is wood, though it is found throughout a plant. The term was introduced by Carl Nägeli in 1858.

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.

Dicotyledon group of plants

The dicotyledons, also known as dicots, are one of the two groups into which all the flowering plants or angiosperms were formerly divided. The name refers to one of the typical characteristics of the group, namely that the seed has two embryonic leaves or cotyledons. There are around 200,000 species within this group. The other group of flowering plants were called monocotyledons or monocots, typically having one cotyledon. Historically, these two groups formed the two divisions of the flowering plants.

Meristem Type of tissue

A meristem is the tissue in most plants containing undifferentiated cells, found in zones of the plant where growth can take place. Meristematic cells give rise to various organs of a plant and are responsible for growth.

A season is a division of the year marked by changes in weather, ecology, and amount of daylight. On Earth, seasons result from Earth's orbit around the Sun and Earth's axial tilt relative to the ecliptic plane. In temperate and polar regions, the seasons are marked by changes in the intensity of sunlight that reaches the Earth's surface, variations of which may cause animals to undergo hibernation or to migrate, and plants to be dormant. Various cultures define the number and nature of seasons based on regional variations.

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.

Monocotyledon important clade of plants

Monocotyledons, commonly referred to as monocots, are flowering plants (angiosperms) the seeds of which typically contain only one embryonic leaf, or cotyledon. They constitute one of the major groups into which the flowering plants have traditionally been divided, the rest of the flowering plants having two cotyledons and therefore classified as dicotyledons, or dicots. However, molecular phylogenetic research has shown that while the monocots form a monophyletic group or clade, the dicotyledons do not. Monocotyledons have almost always been recognized as a group, but with various taxonomic ranks and under several different names. The APG III system of 2009 recognises a clade called "monocots" but does not assign it to a taxonomic rank.

Reproductive anatomy

A collection of flowers forming an inflorescence. Cimicifuga ramosa - spikes.jpg
A collection of flowers forming an inflorescence.

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 fruit containing 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 ovule and 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

History of classification

From 1736, an illustration of Linnaean classification Ehret-Methodus Plantarum Sexualis.jpg
From 1736, an illustration of Linnaean classification

The botanical term "Angiosperm", from the Ancient Greek αγγείον, angeíon (bottle, vessel) and σπέρμα, (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.

An auxanometer, a device for measuring increase or rate of growth in plants NSRW Auxanometer.png
An auxanometer, a device for measuring increase or rate of growth in 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

Monocot (left) and dicot seedlings Monocot vs dicot crop Pengo.jpg
Monocot (left) and dicot seedlings

There are eight groups of living angiosperms:

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]

angiosperms

Amborella

Nymphaeales

Austrobaileyales

magnoliids

Chloranthales

monocots

Ceratophyllum

eudicots

1. Phylogeny of the flowering plants, as of APG III (2009). [13]

angiosperms

Amborella

Nymphaeales

Austrobaileyales

monocots

Chloranthales

magnoliids

Ceratophyllum

eudicots

2. Example of alternative phylogeny (2010) [20]

angiosperms

Amborellales

Nymphaeales

Austrobaileyales

magnoliids

Chloranthales

monocots

Ceratophyllales  

eudicots

basal angiosperms
core angiosperms

3. APG IV (2016) [1]

Evolution

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.

Flowers of Malus sylvestris (crab apple) Malus sylvestris inflorescence, Vosseslag, Belgium.jpg
Flowers of Malus sylvestris (crab apple)

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.[ citation needed ] The fossil plant species Nanjinganthus dendrostyla from Early Jurassic China shares many exclusively angiosperm features, such as a thickened receptacle with ovules, although it is unknown whether it is a crown-group angiosperm or a stem-group angiosperm. [24] [25]

The evolution of seed plants and later angiosperms appears to be the result of two distinct rounds of whole genome duplication events. [26] 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. [27] That event was studied by sequencing the genome of an ancient flowering plant, Amborella trichopoda , [28] and directly addresses Darwin's "abominable mystery."

Flowers and leaves of Oxalis pes-caprae (Bermuda buttercup) Oxalis-pes-caprae-36-Zachi-Evenor.jpg
Flowers and leaves of Oxalis pes-caprae (Bermuda buttercup)

The earliest known macrofossil confidently identified as an angiosperm, Archaefructus liaoningensis , is dated to about 125 million years BP (the Cretaceous period), [29] whereas pollen considered to be of angiosperm origin takes the fossil record back to about 130 million years BP [30] . 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. [31] In 2018, scientists reported that the earliest flowers began about 180 million years ago, 50 million years earlier than thought earlier. [32] 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. [33] [34] 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. [35]

Recent DNA analysis based on molecular systematics [36] [37] 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 [38] 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. [39]

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. [40] 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. [41] The radiation of herbaceous angiosperms occurred much later. [42] 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. [43]

Two bees on a flower head of Creeping Thistle, Cirsium arvense Cirsium arvense with Bees Richard Bartz.jpg
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 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. [44]

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. [45] 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. [46]

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. [47]

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. [48] [49]

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. [50]

Diversity

A poster of twelve different species of flowers of the Asteraceae family Asteracea poster 3.jpg
A poster of twelve different species of flowers of the Asteraceae family
Lupinus pilosus Lupinus-pilosus-2015-Zachi-Evenor-cropped01.jpg
Lupinus pilosus
Bud of a pink rose Rose bud.jpg
Bud of a pink rose

The number of species of flowering plants is estimated to be in the range of 250,000 to 400,000. [51] [52] [53] This compares to around 12,000 species of moss [54] or 11,000 species of pteridophytes, [55] showing that the flowering plants are much more diverse. The number of families in 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] [56]

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, [57] in their APG circumscriptions, are

  1. Asteraceae or Compositae (daisy family): 22,750 species;
  2. Orchidaceae (orchid family): 21,950;
  3. Fabaceae or Leguminosae (bean family): 19,400;
  4. Rubiaceae (madder family): 13,150; [58]
  5. Poaceae or Gramineae (grass family): 10,035;
  6. Lamiaceae or Labiatae (mint family): 7,175;
  7. Euphorbiaceae (spurge family): 5,735;
  8. Melastomataceae or Melastomaceae (melastome family): 5,005;
  9. Myrtaceae (myrtle family): 4,625;
  10. Apocynaceae (dogbane family): 4,555;
  11. Cyperaceae (sedge family): 4,350;
  12. Malvaceae (mallow family): 4,225;
  13. Araceae (arum family): 4,025;
  14. Ericaceae (heath family): 3,995;
  15. Gesneriaceae (gesneriad family): 3,870;
  16. Apiaceae or Umbelliferae (parsley family): 3,780;
  17. Brassicaceae or Cruciferae (cabbage family): 3,710:
  18. Piperaceae (pepper family): 3,600;
  19. Bromeliaceae (bromeliad family): 3,540;
  20. Acanthaceae (acanthus family): 3,500;
  21. Rosaceae (rose family): 2,830;
  22. Boraginaceae (borage family): 2,740;
  23. Urticaceae (nettle family): 2,625;
  24. Ranunculaceae (buttercup family): 2,525;
  25. Lauraceae (laurel family): 2,500;
  26. Solanaceae (nightshade family): 2,460;
  27. Campanulaceae (bellflower family): 2,380;
  28. Arecaceae (palm family): 2,361;
  29. Annonaceae (custard apple family): 2,220;
  30. Caryophyllaceae (pink family): 2,200;
  31. Orobanchaceae (broomrape family): 2,060;
  32. Amaranthaceae (amaranth family): 2,050;
  33. Iridaceae (iris family): 2,025;
  34. Aizoaceae or Ficoidaceae (ice plant family): 2,020;
  35. Rutaceae (rue family): 1,815;
  36. Phyllanthaceae (phyllanthus family): 1,745;
  37. Scrophulariaceae (figwort family): 1,700;
  38. Gentianaceae (gentian family): 1,650;
  39. Convolvulaceae (bindweed family): 1,600;
  40. Proteaceae (protea family): 1,600;
  41. Sapindaceae (soapberry family): 1,580;
  42. Cactaceae (cactus family): 1,500;
  43. 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

Fertilization and embryogenesis

Angiosperm life cycle Angiosperm life cycle diagram-en.svg
Angiosperm life cycle

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

The fruit of the Aesculus or Horse Chestnut tree Aesculus hippocastanum fruit.jpg
The fruit of the Aesculus or Horse Chestnut tree

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. [59]

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

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. [60] 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 [61] and others emphasize DNA repair. [62]

Apomixis

Apomixis (reproduction via asexually formed seeds) is found naturally in about 2.2% of angiosperm genera. [63] One type of apomixis, gametophytic apomixis found in a dandelion species [64] 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

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. [65]

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. [66]

See also

Notes

  1. The major exception to the dominance of terrestrial ecosystems by flowering plants is the coniferous forest.

Related Research Articles

Magnoliales order of plants

The Magnoliales comprise an order of flowering plants.

Canellales botanical name for an order of flowering plants

Canellales is the botanical name for an order of flowering plants, one of the four orders of the magnoliids. It is recognized by the most recent classification of flowering plants, the APG IV system. It is defined to contain two families: Canellaceae and Winteraceae, which comprise 136 species of fragrant trees and shrubs. The Canellaceae are found in tropical America and Africa, and the Winteraceae are part of the Antarctic flora. Although the order was defined based on phylogenetic studies, a number of possible synapomorphies have been suggested, relating to the pollen tube, the seeds, the thickness of the integument, and other aspects of the morphology.

Fertilisation union of gametes of opposite sexes during the process of sexual reproduction to form a zygot

Fertilisation or fertilization, also known as generative fertilisation, insemination, pollination, fecundation, syngamy and impregnation, is the fusion of gametes to initiate the development of a new individual organism or offspring. This cycle of fertilisation and development of new individuals is called sexual reproduction. During double fertilisation in angiosperms the haploid male gamete combines with two haploid polar nuclei to form a triploid primary endosperm nucleus by the process of vegetative fertilisation.

Apomixis replacement of the normal sexual reproduction by asexual reproduction, without fertilization

In botany, apomixis was defined by Hans Winkler as replacement of the normal sexual reproduction by asexual reproduction, without fertilization. Its etymology is Greek for "away from" + "mixing". This definition notably does not mention meiosis. Thus "normal asexual reproduction" of plants, such as propagation from cuttings or leaves, has never been considered to be apomixis, but replacement of the seed by a plantlet or replacement of the flower by bulbils were categorized as types of apomixis. Apomictically produced offspring are genetically identical to the parent plant.

Aristolochiaceae family of plants

The Aristolochiaceae are a family, the birthwort family, of flowering plants with seven genera and about 400 known species belonging to the order Piperales. The type genus is Aristolochia L.

Crossosomatales order of plants

The Crossosomatales are an order, first recognized as such by APG II. They are flowering plants included within the Rosid eudicots.

Buxales order of plants

The Buxales are a small order of eudicot flowering plants, recognized by the APG IV system of 2016. The order includes the family Buxaceae; the families Didymelaceae and Haptanthaceae may also be recognized or may be included in the Buxaceae. Many members of the order are evergreen shrubs or trees, although some are herbaceous perennials. They have separate "male" (staminate) and "female" (carpellate) flowers, mostly on the same plant. Some species are of economic importance either for the wood they produce or as ornamental plants.

<i>Amborella</i> Species of plant

Amborella is a monotypic genus of understory shrubs or small trees endemic to the main island, Grande Terre, of New Caledonia. The genus is the only member of the family Amborellaceae and the order Amborellales and contains a single species, Amborella trichopoda. Amborella is of great interest to plant systematists because molecular phylogenetic analyses consistently place it as the sister group of the remaining flowering plants.

Gymnosperm group of plants, at a varying rank

The gymnosperms, also known as Acrogymnospermae, are a group of seed-producing plants that includes conifers, cycads, Ginkgo, and gnetophytes. The term "gymnosperm" comes from the composite word in Greek: γυμνόσπερμος, literally meaning "naked seeds". The name is based on the unenclosed condition of their seeds. The non-encased condition of their seeds contrasts with the seeds and ovules of flowering plants (angiosperms), which are enclosed within an ovary. Gymnosperm seeds develop either on the surface of scales or leaves, which are often modified to form cones, or solitary as in Yew, Torreya, Ginkgo.

Ovule plant structure and its female reproductive cells

In seed plants, the ovule is the structure that gives rise to and contains the female reproductive cells. It consists of three parts: The integument, forming its outer layer, the nucellus, and the female gametophyte in its center. The female gametophyte — specifically termed a megagametophyte— is also called the embryo sac in angiosperms. The megagametophyte produces an egg cell for the purpose of fertilization.

Endosperm tissue produced inside the seeds of most flowering plants

The endosperm is a tissue produced inside the seeds of most of the flowering plants following fertilization. It is triploid in most species. It surrounds the embryo and provides nutrition in the form of starch, though it can also contain oils and protein. This can make endosperm a source of nutrition in animal diet. For example, wheat endosperm is ground into flour for bread, while barley endosperm is the main source of sugars for beer production. Other examples of endosperm that forms the bulk of the edible portion are coconut "meat" and coconut "water", and corn. Some plants, such as orchids, lack endosperm in their seeds.

Eudicots Clade of flowering plants

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, 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.

Double fertilization complex fertilization mechanism of flowering plants

Double fertilization is a complex fertilization mechanism of flowering plants (angiosperms). This process involves the joining of a female gametophyte with two male gametes (sperm). It begins when a pollen grain adheres to the stigma of the carpel, the female reproductive structure of a flower. The pollen grain then takes in moisture and begins to germinate, forming a pollen tube that extends down toward the ovary through the style. The tip of the pollen tube then enters the ovary and penetrates through the micropyle opening in the ovule. The pollen tube proceeds to release the two sperm in the megagametophyte.

Evolutionary developmental biology (evo-devo) is the study of developmental programs and patterns from an evolutionary perspective. It seeks to understand the various influences shaping the form and nature of life on the planet. Evo-devo arose as a separate branch of science rather recently. An early sign of this occurred in 1999.

Basal angiosperms group of plants

The basal angiosperms are the flowering plants which diverged from the lineage leading to most flowering plants. In particular, the most basal angiosperms were called the ANITA grade which is made up of Amborella, Nymphaeales and Austrobaileyales.

Mesangiospermae clade of plants

Mesangiospermae is a group of flowering plants (angiosperms), informally called "mesangiosperms". They are one of two main clades of angiosperms. It is a name created under the rules of the PhyloCode system of phylogenetic nomenclature. There are about 350,000 species of mesangiosperms. The mesangiosperms contain about 99.95% of the flowering plants, assuming that there are about 175 species not in this group and about 350,000 that are. While such a clade with a similar circumscription exists in the APG III system, it was not given a name.

Superrosids clade of plants

The superrosids are members of a large clade of flowering plants, containing more than 88,000 species, more than a quarter of all angiosperms.

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