Buzz pollination
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Buzz pollination or sonication is a technique used by some bees, such as solitary bees and
bumblebee A bumblebee (or bumble bee, bumble-bee, or humble-bee) is any of over 250 species in the genus ''Bombus'', part of Apidae, one of the bee families. This genus is the only Extant taxon, extant group in the tribe Bombini, though a few extinct r ...
s, to release
pollen Pollen is a powdery substance produced by most types of flowers of seed plants for the purpose of sexual reproduction. It consists of pollen grains (highly reduced Gametophyte#Heterospory, microgametophytes), which produce male gametes (sperm ...
which is more or less firmly held by the
anthers The stamen (: stamina or stamens) is a part consisting of the male reproductive organs of a flower. Collectively, the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filamen ...
. The anthers of buzz-pollinated plant species are typically tubular, with an opening at only one end, and the pollen inside is smooth-grained and firmly attached. With self-fertile plants such as tomatoes, wind may be sufficient to shake loose the pollen through pores in the anther and accomplish
pollination Pollination is the transfer of pollen from an anther of a plant to the stigma (botany), stigma of a plant, later enabling fertilisation and the production of seeds. Pollinating agents can be animals such as insects, for example bees, beetles or bu ...
. Visits by bees may also shake loose some pollen, but more efficient pollination of those plants is accomplished by a few insect species who specialize in
sonication image:Sonicator.jpg, A sonicator at the Weizmann Institute of Science during sonicationSonication is the act of applying sound energy to agitate particles in a sample, for various purposes such as the extraction of multiple compounds from plants, ...
or buzz pollination. In order to release the pollen, solitary bees are able to grab onto the flower and move their flight muscles rapidly, causing the flower and anthers to vibrate, dislodging pollen. Pollination involving vibrations is called buzz pollination.
Honeybee A honey bee (also spelled honeybee) is a eusocial flying insect within the genus ''Apis'' of the bee clade, all native to mainland Afro-Eurasia. After bees spread naturally throughout Africa and Eurasia, humans became responsible for the c ...
s cannot perform buzz pollination. About 9% of the flowers of the world are primarily pollinated using buzz pollination.


Flower morphology

Plants that rely on buzz
pollination Pollination is the transfer of pollen from an anther of a plant to the stigma (botany), stigma of a plant, later enabling fertilisation and the production of seeds. Pollinating agents can be animals such as insects, for example bees, beetles or bu ...
have a unique
anther The stamen (: stamina or stamens) is a part consisting of the male reproductive organs of a flower. Collectively, the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filament ...
shape compared to other flora. In buzz pollinated plants, this process can only happen if pollinators visit the flowers to extract pollen. Only a few insect species are able to pollinate these plants. The flower morphology of buzz pollinated plants is different from that of other flora that do not use this type of pollination. The anthers are completely sealed except for a small pore at the top or have very small slits that open along the sides. The pores and slits are small enough that insects cannot easily enter the anther, but large enough pollen can exit. Because of this shape, they are often referred to as poricidal anthers. These poricidal anthers are only able to release pollen when vibrated at a specific
frequency Frequency is the number of occurrences of a repeating event per unit of time. Frequency is an important parameter used in science and engineering to specify the rate of oscillatory and vibratory phenomena, such as mechanical vibrations, audio ...
. The stigmas of these flowers are often located below the anthers. This is could be an evolutionary strategy to prevent
self-fertilization Autogamy or self-fertilization refers to the Cell fusion, fusion of two gametes that come from one individual. Autogamy is predominantly observed in the form of self-pollination, a Reproduction, reproductive mechanism employed by many flowering pl ...
, also known as selfing, by creating distance between the stigma.


Plants pollinated by buzz pollination

The following plants are pollinated more efficiently by buzz pollination: * All species in ''Primula'' sect. ''Dodecatheon'' (shooting stars) * '' Heliamphora'' * Many members of the family
Solanaceae Solanaceae (), commonly known as the nightshades, is a family of flowering plants in the order Solanales. It contains approximately 2,700 species, several of which are used as agricultural crops, medicinal plants, and ornamental plants. Many me ...
** Most species of the genus ''
Solanum ''Solanum'' is a large and diverse genus of flowering plants, which include three food crops of high economic importance: the potato, the tomato and the eggplant (aubergine, brinjal). It is the largest genus in the nightshade family Solana ...
'' ***
Eggplant Eggplant (American English, US, Canadian English, CA, Australian English, AU, Philippine English, PH), aubergine (British English, UK, Hiberno English, IE, New Zealand English, NZ), brinjal (Indian English, IN, Singapore English, SG, Malays ...
s ***
Potato The potato () is a starchy tuberous vegetable native to the Americas that is consumed as a staple food in many parts of the world. Potatoes are underground stem tubers of the plant ''Solanum tuberosum'', a perennial in the nightshade famil ...
es ***
Tomato The tomato (, ), ''Solanum lycopersicum'', is a plant whose fruit is an edible Berry (botany), berry that is eaten as a vegetable. The tomato is a member of the nightshade family that includes tobacco, potato, and chili peppers. It originate ...
es *** Buffalo bur *** '' Solanum cinereum'', an
Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...
n shrub * '' Hibbertia'' * ''
Dianella Dianella may refer to: * ''Dianella'' (beetle), a species and genus of beetle in the family Carabidae now known as ''Diamella'' * ''Dianella'' (gastropod), a genus of freshwater snails in the family Hydrobiidae * ''Dianella'' (plant), a genus of ...
'' (flax lilies) * Many members of
Ericaceae The Ericaceae () are a Family (biology), family of flowering plants, commonly known as the heath or heather family, found most commonly in acidic and infertile growing conditions. The family is large, with about 4,250 known species spread acros ...
** Some members of the genus ''
Rhododendron ''Rhododendron'' (; : ''rhododendra'') is a very large genus of about 1,024 species of woody plants in the Ericaceae, heath family (Ericaceae). They can be either evergreen or deciduous. Most species are native to eastern Asia and the Himalayan ...
'' ** Some members of the genus ''
Vaccinium ''Vaccinium'' is a common and widespread genus of shrubs or dwarf shrubs in the heath family (Ericaceae). The fruits of many species are eaten by humans and some are of commercial importance, including the cranberry, blueberry, bilberry (wh ...
'' ***
Blueberries Blueberries are a widely distributed and widespread group of perennial flowering plants with blue or purple berries. They are classified in the section ''Cyanococcus'' with the genus ''Vaccinium''. Commercial blueberries—both wild (lowbush) ...
*** Cranberries ** ''
Arctostaphylos ''Arctostaphylos'' (; from "bear" and "bunch of grapes") is a genus of plants comprising the manzanitas () and bearberries. There are about 60 species of ''Arctostaphylos'', ranging from ground-hugging arctic, coastal, and mountain shrub t ...
'' – manzanita * Some
Fabaceae Fabaceae () or Leguminosae,International Code of Nomen ...
** '' Senna''


Examples of buzz pollinating bee species

* '' Amegilla'' sp. * '' Bombus terrestris'' * '' Euglossa sp.'' * ''
Melipona ''Melipona'' is a genus of stingless bees, widespread in warm areas of the Neotropics, from Sinaloa and Tamaulipas (México) to Tucumán Province, Tucumán and Misiones Province, Misiones (Argentina). About 70 species are known.Grüter, C. 2020. ...
sp.'' * '' Nomia sp.'' * '' Oxaea'' sp. * '' Protandrena sp. * '' Xylocopa frontalis''


Evolutionary origins

The earliest evidence of ancestors of plants that use this mode of
pollination Pollination is the transfer of pollen from an anther of a plant to the stigma (botany), stigma of a plant, later enabling fertilisation and the production of seeds. Pollinating agents can be animals such as insects, for example bees, beetles or bu ...
in the fossil record has been dated to the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
period. Also, some extant flora such as members of the family
Myrtaceae Myrtaceae (), the myrtle family, is a family of dicotyledonous plants placed within the order Myrtales. Myrtle, pōhutukawa, bay rum tree, clove, guava, acca (feijoa), allspice, and eucalyptus are some notable members of this group. All ...
show a spectrum of anther shapes including poricidal
anthers The stamen (: stamina or stamens) is a part consisting of the male reproductive organs of a flower. Collectively, the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filamen ...
and are thought to resemble some morphological aspects of ancestral buzz pollinated flowers. As these plants have evolved complex floral structures,
pollinator A pollinator is an animal that moves pollen from the male anther of a flower to the female carpel, stigma of a flower. This helps to bring about fertilization of the ovules in the flower by the male gametes from the pollen grains. Insects are ...
s have coevolved with these plants. Although pollination results from the bees visiting these flowers, this is not the primary reason they visit plants with poricidal anthers. Pollen contains a substantial amount of protein compared to
nectar Nectar is a viscous, sugar-rich liquid produced by Plant, plants in glands called nectaries, either within the flowers with which it attracts pollination, pollinating animals, or by extrafloral nectaries, which provide a nutrient source to an ...
, the sugary liquid the majority of plants produce as a reward for their animal pollinators. Bees eat pollen as well as make a paste with it to feed their
larva A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...
e. The pollen paste is then sealed into the nest to create a reserve for the young bees. Bees rely on this resource for food; therefore they are also dependent on flowers that produce substantial amounts of accessible pollen, including flowers with poricidal anthers. Bees from ''
Bombus A bumblebee (or bumble bee, bumble-bee, or humble-bee) is any of over 250 species in the genus ''Bombus'', part of Apidae, one of the bee families. This genus is the only Extant taxon, extant group in the tribe Bombini, though a few extinct r ...
'' and '' Xylocopa'' are thought to pollinate these flowers because their
adaptive behavior Adaptive behavior is behavior that enables a person (usually used in the context of children) to cope in their environment with greatest success and least conflict with others. This is a term used in the areas of psychology and special education ...
allows them to easily extract pollen that is less available to other insects. Since bees have a source of plentiful pollen that they do not have to compete with other insects for, they are more likely to visit these flowers. This then allows the flowers to be more successful reproductively because the plants maximize their pollen dispersal with each bee visit, and less pollen is lost. The relationship between buzz pollinated plants and bees benefits both groups and could be why poricidal anthers have been successful evolutionarily. Pollinator and flower relationships have been observed in '' Orphium frutescens'', a small shrub that has poricidal anthers. Bees visited these plants outside of the University of Cape Town and continued to visit the plants even when all of the pollen had been extracted. Although the bees did not know the ''O. frutescens'' would benefit from these multiple visits as the plants continue to produce pollen during the flowering season.


Economic impacts

Scientists can now import pollinators, such as bumblebees, where there might be a shortage of pollinators. In New Zealand, the
red clover ''Trifolium pratense'' (from Latin prātum, meaning meadow), red clover, is a herbaceous plant, herbaceous species of flowering plant in the bean family, Fabaceae. It is native to the Old World, but planted and naturalised in many other regions ...
plant population was restored when bumblebees were imported from Europe to help with the pollination. Maintaining red clover population in New Zealand at the time was critical to the country's crop production rate. However, in North America, the managed honeybee industry experienced decline in the early 2000s due to
colony collapse disorder Colony collapse disorder (CCD) is an abnormal phenomenon that occurs when the majority of worker bees in a honey bee bee colony, colony disappear, leaving behind a queen bee, queen, plenty of food, and a few nurse bees to care for the remaining ...
. This has led to higher expenses for farmers from the east to the west coast of the United States and Canada to help pollinate almonds, cucumbers and certain seed plants. One of the most common plants that are assisted by buzz pollination is the tomato ('' Solanum lycopersicum''). In greenhouses worldwide, up to 50 bumblebee colonies are used per hectare during growing season, bringing in values of approximately €13 million a year for a 40,000 hectare globally. Due to competition between producers and an increase in success rate of rearing, the total costs for the bumblebee colonies have diminished severely. This common practice has led to producers branching out and selling not only bumblebees but other insects who may have mutualistic relationships with plants. The total profit of this industry has been recorded to produce over €111 million a year, with €61 million coming in from bumblebees alone.


Environmental impacts

Some conservation organisations expressed concerns about the potential environmental impacts of introducing bumblebees around the world to aid in pollination. In Australia, as bumblebees are not native, and Australia has a number of widely publicised environmental disasters caused by escaped introduced species (" feral species"), research is under way to adapt the use of the Australian native '' Amegilla cingulata'' (blue banded bees) for the same task.


Alternative means of pollination

Greenhouse A greenhouse is a structure that is designed to regulate the temperature and humidity of the environment inside. There are different types of greenhouses, but they all have large areas covered with transparent materials that let sunlight pass an ...
grown tomatoes are unproductive without aid in pollination. Traditionally, pollination has been done by shaking using electric vibrators (one brand name was "Electric Bee"), however, it has been found to be less expensive in human labor and plant breakage to use bumblebees within the greenhouses. Home growers and amateurs can be seen on YouTube using electric toothbrushes to pollinate their flowers.


References


External links

*
Buzz Pollination
Anne Leonard Lab. * {{cite journal , author1=De Luca, P.A. , author2=Vallejo-Marín, M. , year=2013 , title=What's the 'buzz' about? The ecology and evolutionary significance of buzz-pollination , journal=Current Opinion in Plant Biology , volume=16 , issue=4 , pages=429–435 , doi=10.1016/j.pbi.2013.05.002, pmid=23751734 , bibcode=2013COPB...16..271F * Carl Zimmer

''New York Times'', July 11, 2013. * Sue Rosenthal
Buzz Pollination
''Bay Nature'', June 11, 2008.
Buzz pollination and bee learning
Vallejo-Marin Lab Bees Pollination