The monocots (or monocotyledons) are one of the two major groups of flowering plants (or Angiosperms), the other being the dicots (or dicotyledons). In order to reproduce they utilize various strategies such as employing forms of asexual reproduction, restricting which individuals they are sexually compatible with, or influencing how they are pollinated. Nearly all reproductive strategies that evolved in the dicots have independently evolved in monocots as well. [1] Despite these similarities and their close relatedness, monocots and dicots have distinct traits in their reproductive biologies.
Most monocots reproduce sexually through use of seeds that have a single cotyledon, however a great number of monocots reproduce asexually through clonal propagation. Breeding systems that utilize self-incompatibility are much more common than those that utilize self-compatibility. The majority of monocots are animal pollinated (zoophilous), [1] of which most are pollinator generalists. [2] Monocots have mechanisms to promote or suppress cross-fertilization (allogamy) and self-fertilization (autogamy or geitonogamy). The pollination syndromes of monocots can be quite distinct; they include having flower parts in multiples of three, adaptations to pollination by water (hydrogamy), and pollination by sexual deception in orchids. [1]
Reproducing through seeds is the most widespread method of reproduction in both monocots and dicots. However, internal seed structure is vastly different between these groups. The cotyledon is the embryonic leaf within a seed; monocots have one whereas dicots have two. The evolution of having one or two cotyledons may have arisen 200-150 Mya when monocots and dicots are thought to have diverged. [3] [4] Furthermore, the cotyledons in dicot seeds contain the endosperm which acts as the seed’s food storage, while in monocot the endosperm is separated from the cotyledon. [1] Reproduction through seeds is normally a sexual mode of reproduction, however in some cases individuals can asexually produce fertile seeds without pollination, termed apomixis.
Some monocots can reproduce asexually without the need for seeds. Clonal propagation is the production or division of vegetative structures which develop into new individuals that are genetically identical to their progenitor. These vegetative structures can also form enlarged tubers that function as food storage. Monocots constitute the majority of plants with such structures, mainly in the families: Iridaceae, Liliaceae and Amaryllidaceae. There are many different types of clonal propagation, which are classified by the type of tissue propagating.
Monocots can be classified as perfect (having bisexual flowers), monoecious (having separate male and female flowers on the same plant), dioecious (having flowers of only one sex on an individual) and polygamous (having bisexual flowers with male and/or female flowers on the same plant). [1] Plants that are dioecious have no other option but to mate with different individuals, but in all other cases there is the possibility that an individual's pollen may make contact with its own stigma. For this reason, most plants have genetic mechanisms to prevent fertilization from pollen grains that are too closely related to the stigma (self-incompatibility). The mechanisms of breeding systems occur at the molecular level through a biochemical reaction on the stigma that recognizes genetic differences in pollen grains. Depending on the species, individual plants can self-pollinate, individuals plants can cross-pollinate intraspecifically (between individuals of the same species), or individuals can cross-pollinate interspecifically (between individuals of different species) and hybridize. Orchids are known to have weak barriers to hybridization. [1]
Mating with individuals that are too closely related (i.e. with self) may result in inbreeding depression, so it is usually considered advantageous to cross-pollinate intraspecifically, in which case self-incompatibility is utilized. At least 27 families of monocots have genetic mechanisms to ensure self-incompatibility (SI). [5] The most widespread form of self-incompatibility in monocots is gametophytic, [6] meaning compatibility is determined by the genotype of the pollen grain. There are two described mechanisms of gametophytic self-incompatibility that have been shown to occur in four families of dicots (RNase and S-glycoprotein) but none have been found in monocots. [7] [8] However, there is evidence that orchids have an alternative undescribed mechanism of gametophytic self-incompatibility. [7]
Homomorphic sporophytic self-incompatibility has not yet been discovered in monocots. [6] In this form compatibility is determined by the genotype of the anther from which the pollen grain was created. Heteromorphic sporophytic self-incompatibility, a mechanism in heterostylous flowers, has been shown to occur in only one family of monocots, Pontederiaceae. [9] Late-acting (ovarian) self-incompatibility has been described in Agavaceae, Iridaceae, and Amaryllidaceae. [6]
Grasses have a mechanism of self-incompatibility unique to themselves; they employ two unlinked loci, S and Z. When the alleles at these loci are equivalent between a pollen grain and a stigma on which it lands then the pollen grain will be rejected. [10]
Self-compatible (SC) pollination systems are less common than self-incompatibile cross-pollination systems in angiosperms. [11] However, when the probability of cross-pollination is too low it can be advantageous to self-pollinate. Self-pollination is known to be favored in some orchids, rices, and Caulokaempferia coenobialis (Zingiberaceae). [12] [13] [14] [15]
Pollination systems in monocots are just as diverse as in dicots. [1] About two thirds of monocots evolved to be zoophilous (animal pollinated). [1] Others are instead water-pollinated or wind-pollinated such as Cyperaceae, Juncaceae, Sparganiaceae, Typhaceae, and most notably Poaceae. [16] These modes evolved to facilitate transfer of the pollen grain onto the stigma. Most zoophilous monocots are pollinator generalists with the most notable exception being the Orchids. [2] Monocot pollen grains are monocolpate, meaning they have one groove; outer surfaces called exines are smooth.
All monocots utilize either cross-pollination or self-pollination strategies, as do dicots, but the advantage of either strategy depends on ecological factors such as pollinator abundance and competition. These strategies either promote fertilization with self and suppress fertilization with others resulting in self-pollination, or they suppress fertilization with self and promote fertilization with others resulting in cross-pollination. Strategies also exist to suppress fertilization with other species as reproductive barriers.
Pollination strategies have the same function as breeding systems, however they occur at the ecological level or at the level of floral structure rather than at the molecular level on the stigma through genetic recognition
Self-pollination can be prevented by both physical and temporal mechanisms that have evolved in response to the interactions with pollen vectors; these mechanisms make cross-pollination easier to accomplish by lowering the chances of self-pollination. For example, dichogamy, which is the temporal differentiation in the ripening of sexual organs, is common in monocots with both protogynous and protoandrous flowers. Herkogamy, which is the spatial separation of sexual organs, is also present in many monocots. [1]
Self-pollination can occur with or without the aid of animals. When animal-mediated, sexual organs will be positioned closer spatially and temporally, inverse to the strategies of dichogamy and herkogamy. However, when self-pollination is self-induced by the flower, some unique mechanisms have evolved. In Caulokaempferia coenobialis (Zingiberaceae), pollen is transported via a drop of oil that forms on the anther and slowly slides down to the stigma. [13] In the orchid, Paphiopedilum parishii, anthers liquify and touch the stigma with the help of gravity rather than a pollinator. [14] Another orchid, Holcoglossum amesianum, rotates its anther in circles to transfer pollen into its stigma cavity. [15]
Apomixis is asexual reproduction through seeds and does not require pollination. It is distributed throughout the monocot clade in Poales, Asparagales, Liliales, Dioscoreales, and Alismatales. Thus apomixis may have evolved once in a basal ancestor and has since repeatedly become lost. [17]
Pollination syndromes are floral adaptations in response to pollen vectors, such as the production of nectar.
Flower structure is more uniformly distributed within the monocots. Monocot flowers occur with parts having multiples of three; usually there are three stamen, three petals and three sepals (six tepals), and usually just one stigma. [18] However stamens in twos can be found in Cypripedioideae while single fertile stamens can be found in Philydraceae, Zingiberaceae, and as gynostemium in Orchidacaea. Furthermore, flower structures that evolved to trap insects to accomplish pollination are found in many monocot genera. [1] In relation to flower arrangement alone, plants with perfect flowers should be most likely to self-pollinate while dioecious plants should be most likely to cross-pollinate.
Zoophily, or animal pollination, is a method of pollination which utilizes animals as pollen vectors.
In order for pollen to affix to animal bodies, a tryphine coating is usually present in zoophilous pollen to achieve an adhesive pollen grain. [1]
Visual attractants of monocot flowers mainly come from the coloration of tepals. However, when species with small green tepals are zoophilous other organs can evolve to be visually attractive such as having colored bracts (Araceae, Cyclanthaceae, and some Arecaceae), otherwise attraction is based on scent only. The similar pigments used in monocot and dicot flower coloration have independently evolved. [1]
Many monocots produce scent to attract pollinators but perhaps not as many as those that produce nectar. [1] [19]
Most zoophilous monocots produce nectar as a reward and this nectar is alike to nectar of dicots. [1] [19] [20] [21] Carpellary septal nectaries are common and unique to monocots. Nonseptal nectaries are most often epithelial and positioned on the perigonal nectaries of tepals when occurring in monocots. Also, nectar can be produced in perigonal unicellular hairs, a trait only observed in monocots. [1] Monocots do not have disc nectaries whereas in dicots they are widespread. [22] [23]
Like dicots, some zoophilous monocots do not produce nectar and instead offer pollen as the main reward. A few even offer other rewards: oils to bees, starchy tissue to beetles, sleeping holes to bees, and a perfume which Euglossini male bees will collect and present on their legs during mating displays. [1] Deceptive flowers that do not offer actual rewards are much more widespread in monocots than dicots, with the most common perpetrator being the orchids. Orchids commonly provide empty nectar spurs. [1] One genus, Ophrys , is known for its ability to mimic female bees to such a degree that it fools male bees into pseudocopulating with the “female” and thereby pollinating the flower.
Most wind-pollinated plants do not produce nectar, attractive scents, or petals because they are not adapted to pollination by animal vectors. Grasses are a large wind-pollinated group; their stigmas are often feathery to help catch pollen in the wind.
Monocots account for nearly all hydrophilous or water-pollinated plants. These are monocots that are adapted to use water as a vector and constitute most of the aquatic plants. [1] Depending on the species, pollen can either float on the surface and disperse by wind and water currents towards other surface-floating flowers, or pollen can drift underwater to flowers that are submerged. In the later scenario, pollen is without an exine and stigmas are forked. [24]
A few species of bamboos can grow for more than 120 years without flowering. Then at once flowering can simultaneously occur in groves across the world, termed gregarious or mast flowering. This is possible because the trigger to flower is genetically determined and because multiple forests can develop from the clones of one individual. The cause of the trigger is still unknown and unpredictable. During anthesis, or flowering, pollination is wind-mediated but bee pollination has been observed in at least 6 species. When pollination is zoophilous flowers can be fragrant and attract large numbers of pollinator-collecting bees to congregate around the inflorescence and take advantage of this new and abundant source of pollen. After anthesis massive die-offs of all sister groves occur within three years of each other and can have devastating effects. [25] [26] [27] [28]
The dicotyledons, also known as dicots, are one of the two groups into which all the flowering plants (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, typically each having one cotyledon. Historically, these two groups formed the two divisions of the flowering plants.
Monocotyledons, commonly referred to as monocots, are grass and grass-like 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 have two cotyledons and were classified as dicotyledons, or dicots.
Petals are modified leaves that surround the reproductive parts of flowers. They are often brightly coloured or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the corolla. Petals are usually accompanied by another set of modified leaves called sepals, that collectively form the calyx and lie just beneath the corolla. The calyx and the corolla together make up the perianth, the non-reproductive portion of a flower. When the petals and sepals of a flower are difficult to distinguish, they are collectively called tepals. Examples of plants in which the term tepal is appropriate include genera such as Aloe and Tulipa. Conversely, genera such as Rosa and Phaseolus have well-distinguished sepals and petals. When the undifferentiated tepals resemble petals, they are referred to as "petaloid", as in petaloid monocots, orders of monocots with brightly coloured tepals. Since they include Liliales, an alternative name is lilioid monocots.
Pollination is the transfer of pollen from an anther of a plant to the stigma of a plant, later enabling fertilisation and the production of seeds. Pollinating agents can be animals such as insects, for example beetles or butterflies; birds, and bats; water; wind; and even plants themselves. Pollinating animals travel from plant to plant carrying pollen on their bodies in a vital interaction that allows the transfer of genetic material critical to the reproductive system of most flowering plants. When self-pollination occurs within a closed flower. Pollination often occurs within a species. When pollination occurs between species, it can produce hybrid offspring in nature and in plant breeding work.
Iridaceae is a family of plants in order Asparagales, taking its name from the irises. It has a nearly global distribution, with 69 accepted genera with a total of c. 2500 species. It includes a number of economically important cultivated plants, such as species of Freesia, Gladiolus, and Crocus, as well as the crop saffron.
Self-pollination is a form of pollination in which pollen arrives at the stigma of a flower or at the ovule of the same plant. The term cross-pollination is used for the opposite case, where pollen from one plant moves to a different plant.
Plant reproductive morphology is the study of the physical form and structure of those parts of plants directly or indirectly concerned with sexual reproduction.
Self-incompatibility (SI) is a general name for several genetic mechanisms that prevent self-fertilization in sexually reproducing organisms, and thus encourage outcrossing and allogamy. It is contrasted with separation of sexes among individuals (dioecy), and their various modes of spatial (herkogamy) and temporal (dichogamy) separation.
Heterostyly is a unique form of polymorphism and herkogamy in flowers. In a heterostylous species, two or three morphological types of flowers, termed "morphs", exist in the population. On each individual plant, all flowers share the same morph. The flower morphs differ in the lengths of the pistil and stamens, and these traits are not continuous. The morph phenotype is genetically linked to genes responsible for a unique system of self-incompatibility, termed heteromorphic self-incompatibility, that is, the pollen from a flower on one morph cannot fertilize another flower of the same morph.
Entomophily or insect pollination is a form of pollination whereby pollen of plants, especially but not only of flowering plants, is distributed by insects. Flowers pollinated by insects typically advertise themselves with bright colours, sometimes with conspicuous patterns leading to rewards of pollen and nectar; they may also have an attractive scent which in some cases mimics insect pheromones. Insect pollinators such as bees have adaptations for their role, such as lapping or sucking mouthparts to take in nectar, and in some species also pollen baskets on their hind legs. This required the coevolution of insects and flowering plants in the development of pollination behaviour by the insects and pollination mechanisms by the flowers, benefiting both groups. Both the size and the density of a population are known to affect pollination and subsequent reproductive performance.
The stigma is the receptive tip of a carpel, or of several fused carpels, in the gynoecium of a flower.
A flower, also known as a bloom or blossom, is the reproductive structure found in flowering plants. Flowers consist of a combination of vegetative organs – sepals that enclose and protect the developing flower. These petals attract pollinators, and reproductive organs that produce gametophytes, which in flowering plants produce gametes. The male gametophytes, which produce sperm, are enclosed within pollen grains produced in the anthers. The female gametophytes are contained within the ovules produced in the ovary.
Cymodoceaceae is a family of flowering plants, sometimes known as the "manatee-grass family", which includes only marine species.
Pollination syndromes are suites of flower traits that have evolved in response to natural selection imposed by different pollen vectors, which can be abiotic or biotic, such as birds, bees, flies, and so forth through a process called pollinator-mediated selection. These traits include flower shape, size, colour, odour, reward type and amount, nectar composition, timing of flowering, etc. For example, tubular red flowers with copious nectar often attract birds; foul smelling flowers attract carrion flies or beetles, etc.
Plant reproduction is the production of new offspring in plants, which can be accomplished by sexual or asexual reproduction. Sexual reproduction produces offspring by the fusion of gametes, resulting in offspring genetically different from either parent. Asexual reproduction produces new individuals without the fusion of gametes, resulting in clonal plants that are genetically identical to the parent plant and each other, unless mutations occur.
Sexual selection is described as natural selection arising through preference by one sex for certain characteristics in individuals of the other sex. Sexual selection is a common concept in animal evolution but, with plants, it is often overlooked because many plants are hermaphrodites. Flowering plants show many characteristics that are often sexually selected for. For example, flower symmetry, nectar production, floral structure, and inflorescences are just a few of the many secondary sex characteristics acted upon by sexual selection. Sexual dimorphisms and reproductive organs can also be affected by sexual selection in flowering plants.
Reproductive assurance occurs as plants have mechanisms to assure full seed set through selfing when outcross pollen is limiting. It is assumed that self-pollination is beneficial, in spite of potential fitness costs, when there is insufficient pollinator services or outcross pollen from other individuals to accomplish full seed set.. This phenomenon has been observed since the 19th century, when Darwin observed that self-pollination was common in some plants. Constant pollen limitation may cause the evolution of automatic selfing, also known as autogamy. This occurs in plants such as weeds, and is a form of reproductive assurance. As plants pursue reproductive assurance through self-fertilization, there is an increase in homozygosity, and inbreeding depression, due to genetic load, which results in reduced fitness of selfed offspring. Solely outcrossing plants may not be successful colonizers of new regions due to lack of other plants to outcross with, so colonizing species are expected to have mechanisms of reproductive assurance - an idea first proposed by Herbert G. Baker and referred to as Baker's "law" or "rule". Baker's law predicts that reproductive assurance affects establishment of plants in many contexts, including spread by weedy plants and following long-distance dispersal, such as occurs during island colonization. As plants evolve towards increase self-fertilization, energy is redirected to seed production rather than characteristics that increased outcrossing, such as floral attractants, which is a condition known as the selfing syndrome.
Cryptic self-incompatibility (CSI) is the botanical expression that's used to describe a weakened self-incompatibility (SI) system. CSI is one expression of a mixed mating system in flowering plants. Both SI and CSI are traits that increase the frequency of fertilization of ovules by outcross pollen, as opposed to self-pollen.
The pollination of orchids represents a complex aspect of the biology of this plant family, characterized by intricate flower structures and diverse ecological interactions with pollinator. Notably, the topic has garnered significant scientific interest over time, including the attention of Charles Darwin, who is recognized for his contributions to the theory of evolution by natural selection. In 1862, Darwin published his observations on the essential role of insects in orchid pollination in his work The Fertilization of Orchids. He noted that the various strategies employed by orchids to attract their pollinators are complex.
In botany, floral morphology is the study of the diversity of forms and structures presented by the flower, which, by definition, is a branch of limited growth that bears the modified leaves responsible for reproduction and protection of the gametes, called floral pieces.
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