with pollen collected from a night-blooming cereus]] 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 . Pollinating agents can be animals such as insects, for example bees, 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. 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.
In , after the pollen grain (gametophyte) has landed on the stigma, it germinates and develops a pollen tube which grows down the style until it reaches an ovary. Its two travel down the tube to where the gametophyte(s) containing the female gametes are held within the carpel. After entering an ovule through the micropyle, one male nucleus fuses with the Polar body to produce the endosperm tissues, while the other fuses with the egg cell to produce the embryo. Hence the term: "double fertilisation". This process would result in the production of a seed, made of both nutritious tissues and embryo.
In , the ovule is not contained in a carpel, but exposed on the surface of a dedicated support organ, such as the scale of a cone, so that the penetration of carpel tissue is unnecessary. Details of the process vary according to the division of gymnosperms in question. Two main modes of fertilisation are found in gymnosperms: and Ginkgo have motile sperm that swim directly to the egg inside the ovule, whereas and have sperm that are unable to swim but are conveyed to the egg along a pollen tube.
Pollination research covers various fields, including botany, horticulture, entomology, and ecology. The pollination process as an interaction between flower and pollinator was first addressed in the 18th century by Christian Konrad Sprengel. It is important in horticulture and agriculture, because is dependent on fertilisation: the result of pollination. The study of pollination by insects is known as anthecology. There are also studies in economics that look at the positives and negatives of pollination, focused on bees, and how the process affects the pollinators themselves.
The pollen grains are dispersed by the wind to the female, ovulate cone that is made up of many overlapping scales (sporophylls, and thus megasporophylls), each protecting two ovules, each of which consists of a megasporangium (the nucellus) wrapped in two layers of tissue, the integument and the cupule, that were derived from highly modified branches of ancestral gymnosperms. When a pollen grain lands close enough to the tip of an ovule, it is drawn in through the micropyle ( a pore in the integuments covering the tip of the ovule) often by means of a drop of liquid known as a pollination drop. The pollen enters a pollen chamber close to the nucellus, and there it may wait for a year before it germinates and forms a pollen tube that grows through the wall of the megasporangium (=nucellus) where fertilisation takes place. During this time, the megaspore mother cell divides by meiosis to form four haploid cells, three of which degenerate. The surviving one develops as a megaspore and divides repeatedly to form an immature female gametophyte (egg sac). Two or three archegonia containing an egg then develop inside the gametophyte. Meanwhile, in the spring of the second year two sperm cells are produced by mitosis of the body cell of the male gametophyte. The pollen tube elongates and pierces and grows through the megasporangium wall and delivers the sperm cells to the female gametophyte inside. Fertilisation takes place when the nucleus of one of the sperm cells enters the egg cell in the megagametophyte's archegonium.
In flowering plants, the anthers of the flower produce microspores by meiosis. These undergo mitosis to form male gametophytes, each of which contains two haploid cells. Meanwhile, the ovules produce megaspores by meiosis, further division of these form the female gametophytes, which are very strongly reduced, each consisting only of a few cells, one of which is the egg. When a pollen grain adheres to the stigma of a carpel it germinates, developing a pollen tube that grows through the tissues of the style, entering the ovule through the micropyle. When the tube reaches the egg sac, two sperm cells pass through it into the female gametophyte and fertilisation takes place.
Entomophily, pollination by , often occurs on plants that have developed colored petals and a strong Floral scent to attract insects such as bees, wasps, and occasionally ants (Hymenoptera), (Coleoptera), moths and butterflies (Lepidoptera), and flies (Diptera). The existence of insect pollination dates back to the dinosaur era.
Insect pollinators such as ( Apis spp.),
( Bombus spp.), and Butterfly (e.g., Thymelicus flavus) have been observed to engage in flower constancy, which means they are more likely to transfer pollen to other conspecific plants. This can be beneficial for the pollinators, as flower constancy prevents the loss of pollen during interspecific flights and pollinators from clogging stigmas with pollen of other flower species. It also improves the probability that the pollinator will find productive flowers easily accessible and recognisable by familiar clues. The primary insect pollinators are , mostly , but also including sawfly, , and many species of wasps.
Many flowers attract pollinators by odor. For example, orchid bee species such as Euglossa cordata are attracted to orchids this way, and it has been suggested that some orchid species intoxicate bees during visits which can last up to 90 minutes. However, in general, plants that rely on pollen vectors tend to be adapted to their particular type of vector, for example day-pollinated species tend to be brightly coloured and have little odor, but if they are pollinated largely by birds or specialist mammals, they tend to be larger and have larger nectar rewards than species that are strictly insect-pollinated. Night-blooming flowers have little color, but are often very aromatic. Plants with vertebrate pollinators also tend to spread their rewards over longer periods, having long flowering seasons; their specialist pollinators would be likely to starve if the pollination season were too short.
Some flowers have specialized mechanisms to pollination trap to increase effectiveness, attach pollen to specific body parts (as happens in many orchid and Asclepias speciesOllerton, J. & S. Liede. 1997. Pollination systems in the Asclepiadaceae: a survey and preliminary analysis. Biological Journal of the Linnean Society (1997), 62: 593–610.), or require specialized behaviors or morphology in order to extract pollen or nectar. One such syndrome is "buzz pollination" (or "sonication"), where a bee must vibrate at a certain frequency in order to cause pollen to be released from the anthers.
In zoophily, pollination is performed by vertebrates such as and , particularly, , , , , and . Ornithophily or bird pollination is the pollination of flowering plants by birds. Chiropterophily or bat pollination is the pollination of flowering plants by bats. Plants adapted to use bats or moths as pollinators typically have white petals, strong Floral scent and flower at night, whereas plants that use birds as pollinators tend to produce copious nectar and have red petals.
Mammals are not generally thought of as pollinators, but some rodents, bats and marsupials are significant pollinators and some even specialise in such activities. In South Africa certain species of Protea (in particular Protea humiflora, P. amplexicaulis, P. subulifolia, P. decurrens and P. cordata) are adapted to pollination by rodents (particularly Cape Spiny Mouse, Acomys subspinosus) and ( Elephantulus species). The flowers are borne near the ground, are yeasty smelling, not colourful, and sunbirds reject the nectar with its high xylose content. The mice apparently can digest the xylose and they eat large quantities of the pollen. In Australia pollination by flying, gliding and earthbound mammals has been demonstrated.
Reptile pollinators are known, but they form a minority in most ecological situations. They are most frequent and most ecologically significant in island systems, where insect and sometimes also bird populations may be unstable and less species-rich. Adaptation to a lack of animal food and of predation pressure, might therefore favour reptiles becoming more herbivorous and more inclined to feed on pollen and nectar. Most species of lizards in the families that seem to be significant in pollination seem to carry pollen only incidentally, especially the larger species such as Varanidae and Iguanidae, but especially several species of the Gekkonidae are active pollinators, and so is at least one species of the Lacertidae, Podarcis lilfordi, which pollinates various species, but in particular is the major pollinator of Euphorbia dendroides on various Mediterranean islands.
An estimated 48.7% of plant species are either dioecious or self-incompatible obligate out-crossers. It is also estimated that about 42% of flowering plants have a mixed mating system in nature. In the most common kind of mixed mating system, individual plants produce a single type of flower and fruits may contain self-pollinated, out-crossed or a mixture of progeny types.
Pollination also requires consideration of , the plants that serve as the pollen source for other plants. Some plants are self-compatible ( self-fertile) and can pollinate and fertilize themselves. Other plants have chemical or physical barriers to self-pollination.
In agriculture and horticulture pollination management, a good pollenizer is a plant that provides compatible, viable and plentiful pollen and blooms at the same time as the plant that is to be pollinated or has pollen that can be stored and used when needed to pollinate the desired flowers. Hybridization is effective pollination between flowers of different species, or between different breeding lines or populations. see also Heterosis.
are considered self-fertile because a commercial crop can be produced without cross-pollination, though cross-pollination usually gives a better crop. Apples are considered self-incompatible, because a commercial crop must be cross-pollinated. Many commercial fruit tree varieties are grafting cloning, genetics identical. An orchard block of apples of one variety is genetically a single plant. Many growers now consider this a mistake. One means of correcting this mistake is to graft a limb of an appropriate pollenizer (generally a variety of crabapple) every six trees or so.
provide a good example of the mutualism that exists between and angiosperms. Flowers provide bees with nectar (an energy source) and pollen (a source of protein). When bees go from flower to flower collecting pollen they are also depositing pollen grains onto the flowers, thus pollinating them. While pollen and nectar, in most cases, are the most notable reward attained from flowers, bees also visit flowers for other resources such as oil, fragrance, resin and even waxes. It has been estimated that bees originated with the origin or diversification of Flowering plant. In addition, cases of coevolution between bee species and flowering plants have been illustrated by specialized adaptations. For example, long legs are selected for in Rediviva neliana, a bee that collects oil from Diascia capsularis, which have long spur lengths that are selected for in order to deposit pollen on the oil-collecting bee, which in turn selects for even longer legs in R. neliana and again longer spur length in D. capsularis is selected for, thus, continually driving each other's evolution.
Pollination management is a branch of agriculture that seeks to protect and enhance present pollinators and often involves the culture and addition of pollinators in monoculture situations, such as commercial fruit . The largest managed pollination event in the world is in California almond orchards, where nearly half (about one million hives) of the US are trucked to the almond orchards each spring. New York's apple crop requires about 30,000 hives; Maine's blueberry crop uses about 50,000 hives each year. The US solution to the pollinator shortage, so far, has been for commercial beekeepers to become pollination contractors and to migrate. Just as the combine harvesters follow the wheat harvest from Texas to Manitoba, beekeepers follow the bloom from south to north, to provide pollination for many different crops.
In America, bees are brought to commercial plantings of , squash, , strawberry, and many other crops. Honey bees are not the only managed pollinators: a few other species of bees are also raised as pollinators. The alfalfa leafcutter bee is an important pollinator for alfalfa seed in western United States and Canada. are increasingly raised and used extensively for greenhouse and other crops.
The ecology and financial importance of natural pollination by insects to agriculture crops, improving their quality and quantity, becomes more and more appreciated and has given rise to new financial opportunities. The vicinity of a forest or wild with native pollinators near agricultural crops, such as apples, almonds or coffee can improve their yield by about 20%. The benefits of native pollinators may result in forest owners demanding payment for their contribution in the improved crop results – a simple example of the economic value of ecological services. Farmers can also raise native crops in order to promote native bee pollinator species as shown with the native sweat bees L. vierecki in Delaware and L. leucozonium in southwest Virginia.
The American Institute of Biological Sciences reports that native insect pollination saves the United States Agriculture economy nearly an estimated $3.1 billion annually through natural crop production; pollination produces some $40 billion worth of products annually in the United States alone.
Pollination of food crops has become an environmental issue, due to two trends. The trend to monoculture means that greater concentrations of pollinators are needed at bloom time than ever before, yet the area is forage poor or even deadly to bees for the rest of the season. The other trend is the decline of pollinator populations, due to pesticide misuse and overuse, new diseases and of bees, clearfelling, decline of beekeeping, development, removal of hedges and other habitat from , and public concern about bees. Widespread aerial spraying for due to West Nile fears is causing an acceleration of the loss of pollinators. Changes in land use, harmful pesticides, and advancing climate change threaten wild pollinators, key insect species that increase yields of three-fourths of crop varieties and are critical to growing healthy foods.
In some situations, farmers or horticulturists may aim to restrict natural pollination to only permit breeding with the preferred individuals plants. This may be achieved through the use of .
Factors that contribute to pollinator decline include habitat destruction, pesticide, parasitism/diseases, and climate change. The more destructive forms of human disturbances are land use changes such as fragmentation, selective logging, and the conversion to secondary forest habitat. Defaunation of is also an important driver. These alterations are especially harmful due to the sensitivity of the pollination process of plants. Research on tropical palms found that defaunation has caused a decline in seed dispersal, which causes a decrease in genetic variability in this species. Habitat destruction such as fragmentation and selective logging remove areas that are most optimal for the different types of pollinators, which removes pollinators food resources, nesting sites, and leads to isolation of populations. The effect of pesticides on pollinators has been debated because it is difficult to determine that a single pesticide is the cause as opposed to a mixture or other threats. Whether exposure alone causes damage, or if the duration and potency are also factors is unknown. However, have negative effects, as in the case of that harm bee colonies. Many researchers believe it is the synergistic effects of these factors which are ultimately detrimental to pollinator populations.
In the agriculture industry, climate change is causing a "pollinator crisis". This crisis is affecting the production of crops, and the relating costs, due to a decrease in pollination processes.Maglianesi Sandoz, M.A. (2016). Efectos del cambio climático sobre la polinización y la producción agrícola en América Tropical. Revista Ingeniería, 26(1), 11-20. This disturbance can be phenological or spatial. In the first case, species that normally occur in similar seasons or time cycles, now have different responses to environmental changes and therefore no longer interact. For example, a tree may flower sooner than usual, while the pollinator may reproduce later in the year and therefore the two species no longer coincide in time. Spatial disturbances occur when two species that would normally share the same distribution now respond differently to climate change and are shifting to different regions.Butt N, Seabrook L, Maron M, Law BS, Dawson TP, et al. Cascading effects of climate extremes on vertebrate fauna through changes to low latitude tree flowering and fruiting phenology. Global Change Biology. 2015; 21:3267–3277.Visser ME, Both C. Shifts in phenology due to global climate change: the need for a yardstick. Proceedings of the Royal Society of London B. 2005; 272:2561–2569
Butterfly too have suffered due to these modifications. Butterflies are helpful ecological indicators since they are sensitive to changes within the environment like the season, altitude, and above all, human impact on the environment. Butterfly populations were higher within the natural forest and were lower in open land. The reason for the difference in density is the fact that in open land the butterflies would be exposed to desiccation and predation. These open regions are caused by habitat destruction like logging for timber, livestock grazing, and firewood collection. Due to this destruction, butterfly species' diversity can decrease and it is known that there is a correlation in butterfly diversity and plant diversity.
The structure of plant-pollinator networks may have large consequences for the way in which pollinator communities respond to increasingly harsh conditions. Mathematical models, examining the consequences of this network structure for the stability of pollinator communities suggest that the specific way in which plant-pollinator networks are organized minimizes competition between pollinators and may even lead to strong indirect facilitation between pollinators when conditions are harsh. This means that pollinator species together can survive under harsh conditions. But it also means that pollinator species collapse simultaneously when conditions pass a critical point. This simultaneous collapse occurs, because pollinator species depend on each other when surviving under difficult conditions.
Such a community-wide collapse, involving many pollinator species, can occur suddenly when increasingly harsh conditions pass a critical point and recovery from such a collapse might not be easy. The improvement in conditions needed for pollinators to recover, could be substantially larger than the improvement needed to return to conditions at which the pollinator community collapsed.
Over the past decade, beekeepers across the US have reported that the mortality rate of their bee colonies has stayed constant at about 30% every year, making the deaths an expected cost of business for the beekeepers. While the exact cause of this phenomenon is unknown, according to the US Department of Agriculture Colony Collapse Disorder Progress Report it can be traced to factors such as pollution, pesticides, and pathogens from evidences found in areas of the colonies affected and the colonies themselves. Pollution and pesticides are detrimental to the health of the bees and their colonies as the bees' ability to pollinate and return to their colonies are great greatly compromised. Moreover, California's Central Valley is determined by the World Health Organization as the location of country's worst air pollution. Almond pollinating bees, approximately 60% of the bees in the US as mentioned above, will be mixed with bees from thousands of other hives provided by different beekeepers, making them exponentially susceptible to diseases and mites that any of them could be carrying. The deaths do not stop at commercial honeybees as there is evidence of significant pathogen spillover to other pollinators including wild bumble bees, infecting up to 35-100% of wild bees within 2 km radius of commercial pollination. The negative externality of private pollination services is the decline of biodiversity through the deaths of commercial and wild bees. losing about a third of their workforce every year, beekeepers continue to rent out their bees to almond farms due to the high pay from the almond industry. In 2016, a colony rented out for almond pollination gave beekeepers an income of $165 per colony rented, around three times from average of other crops that use the pollination rental service. However, a recent study published in Oxford Academic's Journal of Economic Entomology found that once the costs for maintaining bees specifically for almond pollination, including overwintering, summer management, and the replacement dying bees are considered, almond pollination is barely or not profitable for average beekeepers.
Methods
Biotic pollination
Abiotic pollination
By wind
By water
By rain
Switching methods
Mechanism
Coevolution
In agriculture
Improving pollination in areas with suboptimal bee densities
Environmental impacts
Examples of affected pollinators
Food security and pollinator decline
Plant–pollinator networks
Economics of commercial honeybee pollination
See also
Notes
Further reading
External links
|
|