Buzz pollination
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Buzz pollination or sonication is a technique used by some bees, such as
solitary bees Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyletic lineage within the superfami ...
to release pollen which is more or less firmly held by the anthers. 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 of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by wind. Pollinating agents can be animals such as insects, birds, a ...
. 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 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, microalgae and seawe ...
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 Afro-Eurasia. After bees spread naturally throughout Africa and Eurasia, humans became responsible for the current cosm ...
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 of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by wind. Pollinating agents can be animals such as insects, birds, a ...
have a unique anther 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. It is also occasionally referred to as ''temporal frequency'' for clarity, and is distinct from ''angular frequency''. Frequency is measured in hertz (Hz) which is eq ...
. 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 fusion of two gametes that come from one individual. Autogamy is predominantly observed in the form of self-pollination, a reproductive mechanism employed by many flowering plants. However, species ...
, 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 The genus ''Heliamphora'' ( or ; Greek: ''helos'' "marsh" and ''amphoreus'' " amphora") contains 23 species of pitcher plants endemic to South America.McPherson, S., A. Wistuba, A. Fleischmann & J. Nerz 2011. '' Sarraceniaceae of South America''. ...
'' * Many members of the family Solanaceae ** 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 Solanaceae ...
'' *** Eggplants ***
Potato The potato is a starchy food, a tuber of the plant ''Solanum tuberosum'' and is a root vegetable native to the Americas. The plant is a perennial in the nightshade family Solanaceae. Wild potato species can be found from the southern Unit ...
es ***
Tomato The tomato is the edible berry of the plant ''Solanum lycopersicum'', commonly known as the tomato plant. The species originated in western South America, Mexico, and Central America. The Mexican Nahuatl word gave rise to the Spanish word ...
es *** Buffalo bur *** ''
Solanum cinereum Solanum cinereum is a species of plant in the family Solanaceae, known by the common name Narrawa burr. It is native to open woodland in south eastern Australia. ''Solanum cinereum'' is a small perennial shrub, either sprawling, or erect to o ...
'', an Australian shrub * ''
Hibbertia ''Hibbertia'', commonly known as guinea flowers, is a genus of flowering plants in the family Dilleniaceae. They are usually shrubs with simple leaves and usually yellow flowers with five sepals and five petals. There are about 400 species, mo ...
'' * ''
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 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 c.4250 known species spread across 124 genera, making it th ...
** Some members of the genus '' Rhododendron'' ** 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 (whort ...
'' *** Blueberries ***
Cranberries Cranberries are a group of evergreen dwarf shrubs or trailing vines in the subgenus ''Oxycoccus'' of the genus ''Vaccinium''. In Britain, cranberry may refer to the native species ''Vaccinium oxycoccos'', while in North America, cranberry ...
** ''
Arctostaphylos ''Arctostaphylos'' (; from "bear" and "bunch of grapes") is a genus of plants comprising the manzanitas () and bearberries. They are shrubs or small trees. There are about 60 species, of ''Arctostaphylos'', ranging from ground-hugging arc ...
'' – manzanita * Some Fabaceae ** '' Senna''


Examples of buzz pollinating bee species

* ''
Bombus terrestris ''Bombus terrestris'', the buff-tailed bumblebee or large earth bumblebee, is one of the most numerous bumblebee species in Europe. It is one of the main species used in greenhouse pollination, and so can be found in many countries and areas wher ...
'' * '' Xylocopa frontalis'' * ''
Euglossa ''Euglossa'' is a genus of orchid bees ( Euglossini). Like all their close relatives, they are native to the Neotropics; an introduced population exists in Florida. They are typically bright metallic blue, green, coppery, or golden. '' Euglossa ...
sp.'' * ''
Melipona ''Melipona'' is a genus of stingless bees, widespread in warm areas of the Neotropics, from Sinaloa and Tamaulipas (México) to Tucumán and Misiones (Argentina). About 70 species are known.Grüter, C. 2020. ''Stingless Bees: Their Behaviour, E ...
sp.'' * ''Protandrena sp.'' * '' Nomia sp.'' * ''Oxea sp.''


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 of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by wind. Pollinating agents can be animals such as insects, birds, a ...
in the fossil record has been dated to the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...
period. Also, some extant flora such as members of the family Myrtaceae show a spectrum of anther shapes including poricidal anthers and are thought to resemble some morphological aspects of ancestral buzz pollinated flowers. As these plants have evolved complex floral structures, pollinators have
coevolved In biology, coevolution occurs when two or more species reciprocally affect each other's evolution through the process of natural selection. The term sometimes is used for two traits in the same species affecting each other's evolution, as well ...
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, 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 (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. ...
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 group in the tribe Bombini, though a few extinct related gener ...
'' and ''
Xylocopa Carpenter bees are species in the genus ''Xylocopa'' of the subfamily Xylocopinae. The genus includes some 500 bees in 31 subgenera. The common name "carpenter bee" derives from their nesting behavior; nearly all species burrow into hard plant ma ...
'' 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'', the red clover, is a herbaceous species of flowering plant in the bean family Fabaceae, native to Europe, Western Asia, and northwest Africa, but planted and naturalized in many other regions. Description Red clove ...
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 colony disappear, leaving behind a queen, plenty of food, and a few nurse bees to care for the remaining immature bees. While s ...
. 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 (''
Lycopersicon esculentum The tomato is the edible berry of the plant ''Solanum lycopersicum'', commonly known as the tomato plant. The species originated in western South America, Mexico, and Central America. The Mexican Nahuatl word gave rise to the Spanish word , ...
''). 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 complain 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 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 * 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