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, where 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
 Contents

 1Process
 2Methods
o 2.1Abiotic

 2.1.1By wind

 2.1.2By water

 2.1.3By rain

 2.1.4Switching methods

o 2.2Biotic

 3Mechanism
 4Coevolution
 5In agriculture
o 5.1Improving pollination in areas with suboptimal bee

densities
 6Environmental impacts
o 6.1Examples of affected pollinators

o 6.2Food security and pollinator decline

Process[edit]
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.[4]
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 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.[5][6]

Methods[edit]
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.[7]

Abiotic[edit]
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.
By wind[edit]
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.[8][9][10] 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.[11]
By water[edit]
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.[12] 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.[12]
By rain[edit]
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.[13] In these plants, excess
rain drains allowing the floating pollen to come in contact with the stigma.[13] In rain pollination in
orchids, the rain allows for 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.[14]
Switching methods[edit]
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.[15]

Melissodes desponsus covered in pollen

Biotic[edit]

Hummingbirds typically feed on red flowers

Main article: Pollinator

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.[16] Between 100,000 and 200,000 species of animal act as pollinators of the
world's 250,000 species of flowering plant.[17] 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.[17]
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.[
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. 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.[19]
Insect pollinators such as honey
bees (Apis spp.),[20] bumblebees (Bombus spp.),[21][22] and butterflies (e.g., Thymelicus flavus)[23] have
been observed to engage in flower constancy, which means they are more likely to transfer pollen to
other conspecific plants.[24][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]

Mechanism[edit]

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

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.[5] 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.[5]
 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 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]

 In agriculture[edit]
 Main article: List of crop plants pollinated by bees
 
 An Andrena bee gathers pollen from the stamens of a rose. The female carpel structure appears rough
and globular to the left.


 Bombus ignitus, a popular commercial pollinator in Japan and China[46]
 The most essential staple food crops on the planet,
like corn, wheat, 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.[citation needed]
 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.
 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]

Environmental impacts[edit]
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 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 the 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]

Examples of affected pollinators[edit]

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.[62] 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.[62] The decrease in populations of the honey bee have been attributed to
pesticides, genetically modified crops, fragmentation, parasites and diseases that have been
introduced.[63] 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.[63]
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.[64]

Food security and pollinator decline[edit]

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.[65] 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.[66] 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.[67] The essential nutrients that are necessary in the
human diet are present in plants that rely on animal pollinators.[67] 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.[66]
 Carbon dioxide laser
From Wikipedia, the free encyclopedia

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A test target bursts into flame upon irradiation by a continuous wave kilowatt-level carbon dioxide laser.

The carbon dioxide laser (CO2 laser) was one of the earliest gas lasers to be developed. It was
invented by Kumar Patel of Bell Labs in 1964,[1] and is still one of the most useful. Carbon
dioxide lasers are the highest-power continuous wave lasers that are currently available. They are
also quite efficient: the ratio of output power to pump power can be as large as 20%. The CO2 laser
produces a beam of infrared light with the principal wavelength bands centering on 9.4 and
10.6 micrometers (μm).

Contents

 1Amplification
 2Construction
 3Applications
o 3.1Industrial (cutting and welding)
o 3.2Medical (soft-tissue surgery)
o 3.3Other
 4See also
 5Notes
 6References
 7External links

Amplification[edit]
The active laser medium (laser gain/amplification medium) is a gas discharge which is air- or water-
cooled, depending on the power being applied. The filling gas within the discharge tube consists of
around 10–20% carbon dioxide (CO
2), around 10–20% nitrogen (N

2), a few percent hydrogen (H

2) and/or xenon (Xe) (usually only used in a sealed tube), and the remainder of the gas

mixture helium (He).[citation needed] The specific proportions vary according to the particular laser.
The population inversion in the laser is achieved by the following sequence: electron impact excites
the {v1(1)} vibrational mode quantum state of the nitrogen. Because nitrogen is a homonuclear
molecule, it cannot lose this energy by photon emission, and its excited vibrational modes are
therefore metastable and relatively long-lived. N
2{v1(1)} and CO

2{v3(1)} being nearly perfectly resonant (total molecular energy differential is within 3 cm when
-1

accounting for N
2 anharmonicity, centrifugal distortion and vibro-rotational interaction, which is more than made up for

by the Maxwell speed distribution of translational-mode energy), N

2 collisionally de-excites by transferring its vibrational mode energy to the CO2 molecule, causing the
carbon dioxide to excite to its {v3(1)} (asymmetric stretch) vibrational mode quantum state. The CO
2 then radiatively emits at either 10.6 μm by dropping to the {v1(1)} (symmetric-stretch) vibrational
[i]

mode, or 9.6 μm by dropping to the {v20(2)} (bending) vibrational mode. The carbon dioxide
[i]

molecules then transition to their {v20(0)} vibrational mode ground state from {v1(1)} or {v20(2)} by
collision with cold helium atoms, thus maintaining population inversion. The resulting hot helium
atoms must be cooled in order to sustain the ability to produce a population inversion in the carbon
dioxide molecules. In sealed lasers, this takes place as the helium atoms strike the walls of the laser
discharge tube. In flow-through lasers, a continuous stream of CO2 and nitrogen is excited by the
plasma discharge and the hot gas mixture is exhausted from the resonator by pumps.
Because the excitation energy of molecular vibrational and rotational mode quantum states are low,
the photons emitted due to transition between these quantum states have comparatively lower
energy, and longer wavelength, than visible and near-infrared light. The 9–12 μm wavelength of
CO2 lasers is useful because it falls into an important window for atmospheric transmission (up to
80% atmospheric transmission at this wavelength), and because many natural and synthetic
materials have strong characteristic absorption in this range.[2]
Laser wavelength can be tuned by altering the isotopic ratio of the carbon and oxygen atoms
comprising the CO
2 molecules in the discharge tube.

Construction[edit]
See also: Laser construction

Because CO2 lasers operate in the infrared, special materials are necessary for their construction.
Typically, the mirrors are silvered, while windows and lenses are made of either germanium or zinc
selenide. For high power applications, gold mirrors and zinc selenide windows and lenses are
preferred. There are also diamond windows and lenses in use. Diamond windows are extremely
expensive, but their high thermal conductivity and hardness make them useful in high-power
applications and in dirty environments. Optical elements made of diamond can even be sand
blasted without losing their optical properties. Historically, lenses and windows were made out of salt
(either sodium chloride or potassium chloride). While the material was inexpensive, the lenses and
windows degraded slowly with exposure to atmospheric moisture.
The most basic form of a CO2 laser consists of a gas discharge (with a mix close to that specified
above) with a total reflector at one end, and an output coupler (a partially reflecting mirror) at the
output end.[3]
The CO2 laser can be constructed to have continuous wave (CW) powers between milliwatts (mW)
and hundreds of kilowatts (kW).[4] It is also very easy to actively Q-switch a CO2 laser by means of a
rotating mirror or an electro-optic switch, giving rise to Q-switched peak powers of up
to gigawatts (GW).[5]
Because the laser transitions are actually on vibration-rotation bands of a linear triatomic molecule,
the rotational structure of the P and R bands can be selected by a tuning element in the laser
cavity. Prisms are not practical as tuning elements because most media that transmit in the mid-
infrared absorb or scatter some of the light, so the frequency tuning element is almost always
a diffraction grating. By rotating the diffraction grating, a particular rotational line of the vibrational
transition can be selected. The finest frequency selection may also be obtained through the use of
an etalon. In practice, together with isotopic substitution, this means that a continuous comb of
frequencies separated by around 1 cm−1 (30 GHz) can be used that extend from 880 to 1090 cm−1.
Such "line-tuneable" carbon dioxide lasers[6] are principally of interest in research applications.
The laser's output wavelength is affected by the particular isotopes contained in the carbon dioxide
molecule, with heavier isotopes causing longer wavelength emission. CO
2 lasers can be made to emit from 8.98 to 10.2 μm by selecting the appropriate gas. The table below

shows the output range for nine possible isotope combinations:[2]

Carbon atom First oxygen atom Second oxygen atom

Wavelength (μm)
isotope isotope isotope

C
14 O
16 O
16 9.8–10.2

C
13 O
16 O
16 9.5–9.8

C
12 O
16 O
16 9.1–9.3

C
12 O
16 O
18 9.0–9.2

C
13 O
16 O
18 9.5–9.8

C
12 O
17 O
17 9.0–9.3

C
12 O
18 O
18 9.0–9.2

C
13 O
18 O
18 9.4–9.8
 Carbon atom First oxygen atom Second oxygen atom
Wavelength (μm)
isotope isotope isotope

C
14 18O O
18 9.9–10.2

Applications[edit]

A medical CO2 laser

Industrial (cutting and welding)[edit]

Because of the high power levels available (combined with reasonable cost for the laser), CO2 lasers
are frequently used in industrial applications for cutting and welding, while lower power level lasers
are used for engraving.[7] It is also used in the additive manufacturing process of Selective laser
sintering (SLS).
Medical (soft-tissue surgery)[edit]
Carbon dioxide lasers have become useful in surgical procedures because water (which makes up
most biological tissue) absorbs this frequency of light very well. Some examples of medical uses
are laser surgery and skin resurfacing ("laser facelifts", which essentially consist of vaporizing the
skin to promote collagen formation).[8] CO2 lasers may be used to treat certain skin conditions such
as hirsuties papillaris genitalis by removing bumps or podules. CO2 lasers can be used to remove
vocal fold lesions,[9] such as vocal fold cysts. Researchers in Israel are experimenting with using
CO2 lasers to weld human tissue, as an alternative to traditional sutures.[10]
The 10.6 μm CO2 laser remains the best surgical laser for the soft tissue where both cutting
and hemostasis are achieved photo-thermally (radiantly).[11][12][13][14] CO2 lasers can be used in place of
a scalpel for most procedures, and are even used in places a scalpel would not be used, in delicate
areas where mechanical trauma could damage the surgical site. CO2 lasers are the best suited
for soft tissue procedures in human and animal specialties, as compared to other laser wavelengths.
Advantages include less bleeding, shorter surgery time, less risk of infection, and less post-op
swelling. Applications include gynecology, dentistry, oral and maxillofacial surgery, and many others.
The CO2 laser at the 9.25 - 9.6 μm wavelength is sometimes used in dentistry for hard-tissue
ablation. The hard-tissue is ablated at temperatures as high as 5,000 °C, producing bright thermal
radiation.[15]