Pollination is the act of transferring pollen grains from the male anther of a flower to the female stigma. The goal of every living organism, including plants, is to create offspring for the next generation. One of the ways that plants can produce offspring is by making seeds.
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Pollination
1. Female Xylocopa with pollen collected from night-
blooming cereus
Bee pollinating a plum tree (Prunus
cerasifera).
Pollination
Pollination is the transfer of pollen from a male part of a
plant to a female part of a plant, later enabling fertilisation
and the production of seeds, most often by an animal or
by wind.[1] Pollinating agents are animals such as insects,
birds, and bats; water; wind; and even plants themselves,
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.
In angiosperms, 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 gametes travel down the tube to where the
gametophyte(s) containing the female gametes are held
within the carpel. After entering an ovum cell through the
micropyle, one male nucleus fuses with the polar bodies
to produce the endosperm tissues, while the other fuses with the ovule
to produce the embryo.[2][3] Hence the term: "double fertilization".
This process would result in the production of a seed made of both
nutritious tissues and embryo.
In gymnosperms, 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 fertilization are found in gymnosperms.
Cycads and Ginkgo have motile sperm that swim directly to the egg
inside the ovule, whereas conifers and gnetophytes have sperm that
are unable to swim but are conveyed to the egg along a pollen tube.
The study of pollination spans many disciplines, such as botany,
horticulture, entomology, and ecology. The pollination process as an
interaction between flower and pollen vector was first addressed in
the 18th century by Christian Konrad Sprengel. It is important in
horticulture and agriculture, because fruiting is dependent on
fertilization: the result of pollination. The study of pollination by insects is known as anthecology. There are
also studies in economics that look at the positive and negative benefits of pollination, focused on bees, and
how the process affects the pollinators themselves.
Process of pollination
Methods
Abiotic
By wind
Contents
2. Pollen grains observed in aeroplankton
of South Europe[4]
By water
By rain
Switching methods
Biotic
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 pollination
See also
References
Notes
External links
Pollen germination has three stages; hydration,
activation and pollen tube emergence. The pollen
grain is severely dehydrated so that its mass is
reduced enabling it to be more easily transported
from flower to flower. Germination only takes place
after rehydration, ensuring that premature
germination does not take place in the anther.
Hydration allows the plasma membrane of the
pollen grain to reform into its normal bilayer
organization providing an effective osmotic
membrane. Activation involves the development of
actin filaments throughout the cytoplasm of the cell,
which eventually become concentrated at the point
from which the pollen tube will emerge. Hydration
and activation continue as the pollen tube begins to
grow.[5]
In conifers, the reproductive structures are borne on
cones. The cones are either pollen cones (male) or
ovulate cones (female), but some species are
monoecious and others dioecious. A pollen cone
contains hundreds of microsporangia carried on (or
borne on) reproductive structures called sporophylls. Spore mother cells in the microsporangia divide by
meiosis to form haploid microspores that develop further by two mitotic divisions into immature male
Process of pollination
3. gametophytes (pollen grains). The four resulting cells consist of a large tube cell that forms the pollen tube, a
generative cell that will produce two sperm by mitosis, and two prothallial cells that degenerate. These cells
comprise a very reduced microgametophyte, that is contained within the resistant wall of the pollen grain.[6][7]
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.[7]
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.[6]
Pollination may be biotic or abiotic. Biotic pollination relies on living pollinators to move the pollen from one
flower to another. Abiotic pollination relies on wind, water or even rain. About 80% of angiosperms rely on
biotic pollination.[8]
Abiotic pollination uses nonliving methods such as wind and water to move pollen from one flower to another.
This allows the plant to spend energy directly on pollen rather than on attracting pollinators with flowers and
nectar.
Some 98% of abiotic pollination is anemophily, pollination by wind. This probably arose from insect
pollination, most likely due to changes in the environment or the availability of pollinators.[9][10][11] The
transfer of pollen is more efficient than previously thought; wind pollinated plants have developed to have
specific heights, in addition to specific floral, stamen and stigma positions that promote effective pollen
dispersal and transfer.[12]
Methods
Abiotic
By wind
By water
4. Melissodes desponsus covered in
pollen
Hummingbirds typically feed on red
flowers
A bee on a strawberry blossom
Pollination by water, hydrophily, uses water to transport pollen, sometimes as whole anthers; these can travel
across the surface of the water to carry dry pollen from one flower to another.[13] In Vallisneria spiralis, an
unopened male flower floats to the surface of the water, and, upon reaching the surface, opens up and the
fertile anthers project forward. The female flower, also floating, has its stigma protected from the water, while
its sepals are slightly depressed into the water, allowing the male flowers to tumble in.[13]
Rain pollination is used by a small percentage of plants. Heavy rain discourages insect pollination and
damages unprotected flowers, but can itself disperse pollen of suitably adapted plants, such as Ranunculus
flammula, Narthecium ossifragum, and Caltha palustris.[14] In these plants, excess rain drains allowing the
floating pollen to come in contact with the stigma.[14] In some orchids ombrophily occurs, and rain water
splashes cause the anther cap to be removed, allowing for the pollen to be exposed. After exposure, raindrops
causes the pollen to be shot upward, when the stipe pulls them back, and then fall into the cavity of the stigma.
Thus, for the orchid Acampe rigida, this allows the plant to self-pollinate, which is useful when biotic
pollinators in the environment have decreased.[15]
It is possible for a plant have varying pollination methods, including both biotic and abiotic pollination. The
orchid Oeceoclades maculata uses both rain and butterflies, depending on its environmental conditions.[16]
More commonly, pollination
involves pollinators (also
called pollen vectors):
organisms that carry or move
the pollen grains from the
anther of one flower to the
receptive part of the carpel or
pistil (stigma) of another.[17]
Between 100,000 and
200,000 species of animal act as pollinators of the world's 250,000
species of flowering plant.[18] The majority of these pollinators are
insects, but about 1,500 species of birds and mammals visit flowers
and may transfer pollen between them. Besides birds and bats which
are the most frequent visitors, these include monkeys, lemurs,
squirrels, rodents and possums.[18]
Entomophily, pollination by insects, often occurs on plants that have
developed colored petals and a strong scent to attract insects such as,
bees, wasps and occasionally ants (Hymenoptera), beetles
(Coleoptera), moths and butterflies (Lepidoptera), and flies (Diptera).
The existence of insect pollination dates back to the dinosaur era.[19]
In zoophily, pollination is performed by vertebrates such as birds and bats, particularly, hummingbirds,
sunbirds, spiderhunters, honeyeaters, and fruit bats. Ornithophily or bird pollination is the pollination of
flowering plants by birds. Chiropterophily or bat pollination is the pollination of flowering plants by bats.
By rain
Switching methods
Biotic
Play media
5. Plants adapted to use bats or moths as pollinators typically have white petals, strong scent and flower at night,
whereas plants that use birds as pollinators tend to produce copious nectar and have red petals.[20]
Insect pollinators such as honey bees (Apis spp.),[21] bumblebees (Bombus spp.),[22][23] and butterflies (e.g.,
Thymelicus flavus)[24] have been observed to engage in flower constancy, which means they are more likely
to transfer pollen to other conspecific plants.[25] 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.[26]
Some flowers have specialized mechanisms to trap pollinators to increase effectiveness.[27] Other flowers will
attract pollinators by odor. For example, bee species such as Euglossa cordata are attracted to orchids this way,
and it has been suggested that the bees will become intoxicated during these visits to the orchid flowers, which
last up to 90 minutes.[28] 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, 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. They 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.[27]
As for the types of pollinators, 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.[29] 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.[30]
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)[31] and elephant shrews (Elephantulus
species).[32] 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.[33] In Australia pollination by flying, gliding and earthbound mammals has been demonstrated.[34]
Examples of pollen vectors include many species of wasps, that transport pollen of many plant species, being
potential or even efficient pollinators.[35]
Pollination can be accomplished by cross-pollination or by self-pollination:
Cross-pollination, also called allogamy, occurs when pollen is delivered from the stamen of
one flower to the stigma of a flower on another plant of the same species.[6] Plants adapted for
cross-pollination have several mechanisms to prevent self-pollination; the reproductive organs
may be arranged in such a way that self-fertilisation is unlikely, or the stamens and carpels may
mature at different times.[6]
Self-pollination occurs when pollen from one flower pollinates the same flower or other flowers
of the same individual.[36] It is thought to have evolved under conditions when pollinators were
Mechanism
6. A European honey bee collects
nectar, while pollen collects on its
body.
Africanized honey bees immersed in
Opuntia engelmannii cactus Pollen
Diadasia bee straddles
cactus carpels
not reliable vectors for pollen transport, and is most often
seen in short-lived annual species and plants that colonize
new locations.[37] Self-pollination may include autogamy,
where pollen is transferred to the female part of the same
flower; or geitonogamy, when pollen is transferred to
another flower on the same plant.[38] Plants adapted to
self-fertilize often have similar stamen and carpel lengths.
Plants that can pollinate themselves and produce viable
offspring are called self-fertile. Plants that cannot fertilize
themselves are called self-sterile, a condition which
mandates cross-pollination for the production of
offspring.[38]
Cleistogamy: is self-pollination that occurs before the
flower opens. The pollen is released from the anther within
the flower or the pollen on the anther grows a tube down
the style to the ovules. It is a type of sexual breeding, in
contrast to asexual systems such as apomixis. Some
cleistogamous flowers never open, in contrast to
chasmogamous flowers that open and are then pollinated.
Cleistogamous flowers are by necessity found on self-
compatible or self-fertile plants.[39] Although certain
orchids and grasses are entirely cleistogamous, other
plants resort to this strategy under adverse conditions.
Often there may be a mixture of both cleistogamous and
chasmogamous flowers, sometimes on different parts of
the plant and sometimes in mixed inflorescences. The
ground bean produces cleistogamous flowers below
ground, and mixed cleistogamous and chasmogamous
flowers above.[40]
7. Geranium incanum,
like most geraniums
and pelargoniums,
sheds its anthers,
sometimes its
stamens as well, as
a barrier to self-
pollination. This
young flower is
about to open its
anthers, but has not
yet fully developed
its pistil.
These Geranium
incanum flowers have
opened their anthers,
but not yet their
stigmas. Note the
change of colour that
signals to pollinators
that it is ready for
visits.
This Geranium
incanum flower has
shed its stamens,
and deployed the
tips of its pistil
without accepting
pollen from its own
anthers. (It might of
course still receive
pollen from younger
flowers on the same
plant.)
An estimated 48.7% of plant species are either dioecious or self-incompatible obligate out-crossers.[41] It is
also estimated that about 42% of flowering plants have a mixed mating system in nature.[42] 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 pollenizers, 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.
Peaches 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 grafted clones, genetically
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.
The first fossil record for abiotic pollination is from fern-like plants in the late Carboniferous period.
Gymnosperms show evidence for biotic pollination as early as the Triassic period. Many fossilized pollen
grains show characteristics similar to the biotically dispersed pollen today. Furthermore, the gut contents, wing
structures, and mouthpart morphology of fossilized beetles and flies suggest that they acted as early pollinators.
The association between beetles and angiosperms during the early Cretaceous period led to parallel radiations
Coevolution
8. The wasp Mischocyttarus
rotundicollis transporting pollen
grains of Schinus terebinthifolius
Melittosphex burmensis, the oldest
bee fossil, from the Cretaceous
An Andrena bee gathers pollen from
the stamens of a rose. The female
carpel structure appears rough and
globular to the left.
of angiosperms and insects into the late Cretaceous. The evolution of
nectaries in late Cretaceous flowers signals the beginning of the
mutualism between hymenopterans and angiosperms.
Bees provide a good example of the mutualism that exists between
hymenopterans 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.[43] It has been estimated that bees
originated with the origin or diversification of angiosperms.[44] 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.[45]
The most essential staple food crops on the planet, like wheat, maize,
rice, soybeans and sorghum[47][48] are wind pollinated or self
pollinating. When considering the top 15 crops contributing to the
human diet globally in 2013, slightly over 10% of the total human diet
of plant crops (211 out of 1916 kcal/person/day) is dependent upon
insect pollination.[47]
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 orchards. The largest managed pollination event in the world is in
Californian almond orchards, where nearly half (about one million
hives) of the US honey bees 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 cucumbers, squash, melons, strawberries, 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. Bumblebees are increasingly raised and used extensively for greenhouse tomatoes and other crops.
In agriculture
9. Bombus ignitus, a popular
commercial pollinator in Japan and
China[46]
The ecological and financial importance of natural pollination by
insects to agricultural 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 grasslands 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
L. vierecki in Delaware[49] and L. leucozonium in southwest
Virginia.[50]
The American Institute of Biological Sciences reports that native
insect pollination saves the United States agricultural economy nearly
an estimated $3.1 billion annually through natural crop production;[51]
pollination produces some $40 billion worth of products annually in
the United States alone.[52]
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 parasites of bees, clearcut logging, decline
of beekeeping, suburban development, removal of hedges and other habitat from farms, and public concern
about bees. Widespread aerial spraying for mosquitoes due to West Nile fears is causing an acceleration of the
loss of pollinators.
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 pollination bags.
In some instances growers’ demand for beehives far exceeds the available supply. The number of managed
beehives in the US has steadily declined from close to 6 million after WWII, to less than 2.5 million today. In
contrast, the area dedicated to growing bee-pollinated crops has grown over 300% in the same time period.
Additionally, in the past five years there has been a decline in winter managed beehives, which has reached an
unprecedented rate of colony losses at near 30%.[53][54][55][56] At present, there is an enormous demand for
beehive rentals that cannot always be met. There is a clear need across the agricultural industry for a
management tool to draw pollinators into cultivations and encourage them to preferentially visit and pollinate
the flowering crop. By attracting pollinators like honey bees and increasing their foraging behavior,
particularly in the center of large plots, we can increase grower returns and optimize yield from their plantings.
ISCA Technologies,[57] from Riverside California, created a semiochemical formulation called SPLAT
Bloom, that modifies the behavior of honey bees, inciting them to visit flowers in every portion of the field.
Loss of pollinators, also known as Pollinator decline (of which colony collapse disorder is perhaps the most
well known) has been noticed in recent years. These loss of pollinators have caused a disturbance in early
plant regeneration processes such as seed dispersal and pollination. Early processes of plant regeneration
greatly depend on plant-animal interactions and because these interactions are interrupted, biodiversity and
ecosystem functioning are threatened.[58] Pollination by animals aids in the genetic variability and diversity
Improving pollination in areas with suboptimal bee densities
Environmental impacts
10. within plants because it allows for out-crossing instead for self-crossing. Without this genetic diversity there
would be a lack of traits for natural selection to act on for the survival of the plant species. Seed dispersal is
also important for plant fitness because it allows plants the ability to expand their populations. More than that,
it permits plants to escape environments that have changed and have become difficult to reside in. All of these
factors show the importance of pollinators for plants, which are a significant part of the foundation for a stable
ecosystem. If only a few species of plants depended on Loss of pollinators is especially devastating because
there are so many plant species rely on them. More than 87.5% of angiosperms, over 75% of tropical tree
species, and 30-40% of tree species in temperate regions depend on pollination and seed dispersal.[58]
Factors that contribute to pollinator decline include habitat destruction, pesticide, parasitism/diseases, and
climate change.[59] The more destructive forms of human disturbances are land use changes such as
fragmentation, selective logging, and the conversion to secondary forest habitat.[58] Defaunation of frugivores
is also an important driver.[60] These alterations are especially harmful due to the sensitivity of the pollination
process of plants.[58] Research on tropical palms found that defaunation has caused a decline in seed dispersal,
which causes a decrease in genetic variability in this species.[60] 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.[61] 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.[61] Whether exposure alone causes damage, or if the duration and potency are also
factors is unknown.[61] However, insecticides have negative effects, as in the case of neonicotinoids that harm
bee colonies. Many researchers believe it is the synergistic effects of these factors which are ultimately
detrimental to pollinator populations.[59]
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.[62] 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.[63][64]
The most known and understood pollinator, bees, have been used as the prime example of the decline in
pollinators. Bees are essential in the pollination of agricultural crops and wild plants and are one of the main
insects that perform this task.[65] Out of the bees species, the honey bee or Apis mellifera has been studied the
most and in the United States, there has been a loss of 59% of colonies from 1947 to 2005.[65] The decrease in
populations of the honey bee have been attributed to pesticides, genetically modified crops, fragmentation,
parasites and diseases that have been introduced.[66] There has been a focus on neonicotinoids effects on
honey bee populations. Neonicotinoids insecticides have been used due to its low mammalian toxicity, target
specificity, low application rates, and broad spectrum activity. However, the insecticides are able to make its
way throughout the plant, which includes the pollen and nectar. Due to this, it has been shown to effect on the
nervous system and colony relations in the honey bee populations.[66]
Butterflies 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.[67]
Examples of affected pollinators
11. Besides the imbalance of the ecosystem caused by the decline in pollinators, it may jeopardise food security.
Pollination is necessary for plants to continue their populations and 3/4 of the plant species that contribute to
the world's food supply are plants that require pollinators.[68] Insect pollinators, like bees, are large
contributors to crop production, over 200 billion dollars worth of crop species are pollinated by these
insects.[61] Pollinators are also essential because they improve crop quality and increase genetic diversity,
which is necessary in producing fruit with nutritional value and various flavors.[69] Crops that do not depend
on animals for pollination but on the wind or self-pollination, like corn and potatoes, have doubled in
production and make up a large part of the human diet but do not provide the micronutrients that are
needed.[70] The essential nutrients that are necessary in the human diet are present in plants that rely on animal
pollinators.[70] There have been issues in vitamin and mineral deficiencies and it is believed that if pollinator
populations continue to decrease these deficiencies will become even more prominent.[69]
Wild pollinators often visit a large number of plant species and plants are visited by a large number of
pollinator species. All these relations together form a network of interactions between plants and pollinators.
Surprising similarities were found in the structure of networks consisting out of the interactions between plants
and pollinators. This structure was found to be similar in very different ecosystems on different continents,
consisting of entirely different species.[71]
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[72] and may even lead to strong
indirect facilitation between pollinators when conditions are harsh.[73] 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.[73]
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.[73]
While there are 200,000 - 350,000 different species of animals that help pollination, bees are responsible for
majority of the pollination for consumed crops, providing between $235 and $577 US billion of benefits to
global food production.[74] Since the early 1900s, beekeepers in the United States started renting out their
colonies to farmers to increase the farmer's crop yields, earning additional revenue from providing privatized
pollination. As of 2016, 41% of an average US beekeeper's revenue comes from providing such pollination
service to farmers, making it the biggest proportion of their income, with the rest coming from sales of honey,
beeswax, government subsidy, etc.[75] This is an example of how a positive externality, pollination of crops
from beekeeping and honey-making, was successfully accounted for and incorporated into the overall market
for agriculture. On top of assisting food production, pollination service provide beneficial spillovers as bees
germinate not only the crops, but also other plants around the area that they are set loose to pollinate,
increasing biodiversity for the local ecosystem.[76] There is even further spillover as biodiversity increases
ecosystem resistance for wildlife and crops.[77] Due to their role of pollination in crop production, commercial
Food security and pollinator decline
Plant–pollinator networks
Economics of pollination
12. The graph shows the number of
honeybee colonies in the U.S. from
1982 to 2015,
The graph shows the average dollar
amount per colonies received by
beekeepers depending on the
pollinated crop.
honeybees are considered to be livestock by the US Department of
Agriculture. The impact of pollination varies by crop. For example,
almond production in the United States, an $11 billion industry based
almost exclusively in the state of California, is heavily dependent on
bees for pollination of almond trees. Almond industry uses up to 82%
of the services in the pollination market. Each February, around 60%
of the all bee colonies in the US are moved to California's Central
Valley.[78]
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.[79] 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.[80] Moreover, California's Central Valley is determined by the World Health Organization as the
location of country's worst air pollution.[81] 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.[78] 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.[82] The negative externality of private pollination services is the decline of
biodiversity through the deaths of commercial and wild bees.
Despite 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.[83] 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.[84]
Canadian Pollination Initiative
Cheating (biology)
Floral color change
Fruit tree pollination
Hand-pollination
Paul Knuth
Hermann Müller (botanist)
Plant reproductive morphology
Pollen DNA barcoding
See also
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Notes
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