Pigeons and doves are in the order Columbiformes and family Columbidae. There are five subfamilies within Columbidae, 42 genera and 308 species. They are easily recognizable and have a world-wide distribution (although they are not found in Antarctica). They live in almost all types of terrestrial habitats from desert to dense forest and large urban areas. Pigeons and doves are stocky birds that range from 15 to 75 cm long. Many of the seed-eating columbids are buff, grey and brown colors, while the fruit-eaters are often more brightly colored. Many have ornamentation and iridescent feathers on the neck, breast, back, wings and face. They range from solitary to extremely social; the now extinct passenger pigeons (Ectopistes migratorius) are reported to have occurred in flocks of up to two million birds that were so dense that they blocked out the sun.
Pigeons and doves are a cosmopolitan family (although they are not found in Antarctica or the high arctic). The highest diversity of columbids occurs in South America, Australasia and the Pacific Islands. Some species (for example, Rock Doves (Columba livia)) have been introduced throughout much of their range.
Biogeographic Regions: nearctic (Introduced , Native ); palearctic (Native ); oriental (Native ); ethiopian (Native ); neotropical (Introduced , Native ); australian (Introduced , Native ); oceanic islands (Introduced , Native )
Other Geographic Terms: cosmopolitan ; island endemic
Pigeons and doves are stocky birds that range from 15 to 75 cm long and weigh from 30 to over 2000 g. The smaller species within Columbidae are often called doves and the larger species pigeons, but these names do not necessarily reflect true differences and are often used interchangeably. Columbids have small heads and short beaks and legs. Their flight muscles may make up to 44 percent of the bird’s body weight and allow them to have excellent flying capabilities and maneuverability. Wing-shape is often a good indicator of the species’ migratory behavior. They have soft skin at the base of their bills and a ring of bare skin around their eyes that can be red, blue, yellow or white. Columbids have a bilobed crop that produces “crop-milk” (or "pigeon milk") that they feed their young.
Columbids can be divided in to seed-eating and fruit-eating species. Many of the seed-eating columbids are buff, grey and brown colors while the fruit-eaters are often more brightly colored. Typical pigeons (subfamily Columbinae) are usually grey, brown and/or pink. Fruit-eating pigeons (subfamily Treroninae) are more colorful with oranges and greens. Crowned pigeons (subfamily Gourinae) are grey with pink or chestnut underparts and a white wing patch. Tooth-billed pigeons (subfamily Didunculinae) are chestnut colored on the back and wings and dark green elsewhere. Many doves and pigeons have ornamentation (such as crests and colorful eye rings) and iridescent feathers on the neck, breast, back, wings and face. They range from sexually monomorphic to sexually dimorphic, and molt annually after breeding.
Sexual Dimorphism: sexes alike; male larger; sexes colored or patterned differently; male more colorful; ornamentation
Other Physical Features: endothermic ; bilateral symmetry
Columbids are found in almost all terrestrial habitats from temperate areas to the tropics including: lowland rainforest, highland forest, tropical deciduous forest, riparian forest, boreal forest, savanna, desert, cliff, chaparral, coral atolls, mangroves, swamp forest, woodland edge, agricultural areas, suburban and urban areas. The highest diversity of pigeons and doves occurs in tropical rainforests. They can be found from sea level to 5000 m and their excellent flying abilities have allowed them to colonize oceanic islands.
Habitat Regions: temperate ; tropical ; terrestrial
Terrestrial Biomes: desert or dune ; savanna or grassland ; chaparral ; forest ; rainforest ; scrub forest ; mountains
Aquatic Biomes: coastal
Other Habitat Features: urban ; suburban ; agricultural
Pigeons and doves can be seen feeding in flocks. They are primarily grainivorous and frugivorous, but occasionally they eat insects, snails, worms, lizards, leaves, buds and flowers. Seeds are picked up off the ground and eaten whole and fruits are plucked from trees. Grainivorous species have specialized gizzards, intestines and esophagi that help them eat and digest seeds. Grain is stored in their crops and ground by the grit in their gizzard. Grainivorous species need to drink a lot of water in order to digest seeds. Desert species get their water from succulent plants and have the ability to drink saline water. Columbids drink by submerging their beaks into the water and sucking the water up, they do not scoop water in their beaks and lift their heads to swallow like most birds.
Size differences among columbids often reflect dietary differences and allow for resource partitioning. Larger birds eat larger fruit than smaller birds and smaller birds can feed on thinner branches and reach fruit that the larger birds can not reach.
Primary Diet: herbivore (Frugivore , Granivore )
Pigeons and doves are important seed dispersers and are host to a number of feather parasites (including Columbicola columbae and Campanulotes bidentatus). In addition, they can carry human diseases.
Ecosystem Impact: disperses seeds
Pigeons and doves often form feeding flocks, which allow for increased vigilance and reduce the chance that any one bird will be caught by a predator. They also use broken wing displays to draw predators from the nest. In response to high rates of nest predation, columbids have developed short incubation and nestling periods. Snakes (suborder Serpentes) are common nest predators and falcons (family Falconidae) and other birds of prey (order Falconiformes) feed on adults.
- snakes (Serpentes)
- falcons (Falconidae)
- birds of prey (Falconiformes)
Felis silvestris libyca
Based on studies in:
USA: Arizona, Sonora Desert (Desert or dune)
India, Rajasthan Desert (Desert or dune)
This list may not be complete but is based on published studies.
Known prey organisms
seeds of other plants
Based on studies in:
USA: Arizona, Sonora Desert (Desert or dune)
India, Rajasthan Desert (Desert or dune)
This list may not be complete but is based on published studies.
Life History and Behavior
Pigeons and doves have a variety of songs and calls that they use to find mates, signal danger, and defend territories. Males have special vocalizations that are only used in courtship and advertising. Both males and females sing; most songs are flute-like cooing noises that differ in the length of each note and in the interval between notes. Some species sound like a whistle, and others sound more like a croak. Small columbid species have higher-pitched calls than larger species. They will sometimes call in duets. Some species make quiet purring sounds that function in mate-bonding. Adults will respond to song playbacks. Young birds have begging calls and the results of cross-fostering experiments show that songs are innate and are not learned from their parents.
Pigeons and doves have a variety of courtship displays (see Reproduction: Mating Systems). They also have threat displays in which they spread and raise their wings and spread their tail. If the display does not work to repel and intruder, they will “buffet” with their wings or peck at the intruder.
Columbids are excellent navigators and use both the magnetic field of the planet and the position of the sun to find their way.
Communication Channels: visual ; tactile ; acoustic
Other Communication Modes: duets
Perception Channels: visual ; tactile ; acoustic ; chemical ; magnetic
The oldest recorded columbid is a mourning dove (Zenaida macroura) that lived 19 years and 4 months. Adult annual survival has been estimated to be 40 to 65 percent.
Pigeons and doves are monogamous, and many have the same mate from year to year. They have numerous displays that are performed either on the ground or in the air. For example, while on the ground, males of some species squat, lift their tail, lower their head, twitch their wings and scratch the ground with their feet while calling. Some species have aerial displays that usually involve wing claps (most forest and ground living species do not have aerial displays). Aerial displays are used in courtship and to indicate territory boundaries. During a pre-copulation display, a male inflates his crop, bows, spreads his tail feathers, pirouettes and calls. This display varies among species, and birds with ornaments and colorful plumage usually show them off during the display. Some species of pigeons and doves also engage in courtship feeding and/or mate guarding.
Mating System: monogamous
Breeding is triggered by food availability and photoperiod and can be seasonal or year-round, depending on the species. Some columbids breed colonially, some solitarily. The males bring nest material to the females who build the nest. The nest is a platform or shallow cup of twigs and stems built on a crevice, cliff, tree or the ground. Columbids will re-use nests and will build nests on top of abandoned bird nests. Nest building usually lasts two to four days and nest sites are defended.
Clutch size is usually one to two eggs (occasionally three). Frugivorous species usually have only one egg; fruit is low in protein so the birds can not raise more than one chick. The eggs are white or buff colored and unmarked. Both males and females incubate, but females usually spend more time incubating than males. Species that live in the desert wet their stomach feathers before incubating to help cool the eggs by evaporation. Incubation lasts 11 to 30 days, and hatching can be either synchronous or asynchronous. Chicks are altricial and are fed by both parents. Chicks fledge in 10 to 36 days (earlier if disturbed) and may continue to receive food from their parents for 30 to 40 days.
Breeding pairs can have up to five broods in one breeding season. Their short breeding cycle allows pigeons and doves to have more broods to compensate for their small brood sizes and relatively high rates of predation. Fledglings grow their adult plumage a few months after fledging and reach sexual maturity in 6 to 12 months.
Key Reproductive Features: iteroparous ; seasonal breeding ; year-round breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; fertilization (Internal ); oviparous
Both males and females incubate, but females usually spend more time incubating than males. Species that live in the desert wet their stomach feathers before incubating to help cool the eggs by evaporation. Incubation lasts 11 to 30 days. Chicks are altricial and are fed by both parents. The chicks are usually fed crop-milk for three to four days and are then fed seeds and fruit, however, some continue to be fed crop-milk even after they have fledged. Crop-milk is made in the crop of the adult birds and is 75 to 77 percent water, 11 to 13 percent protein, 5 to 7 percent fat and 1.2 to 1.8 percent minerals and amino acids. Nestling pigeons and doves grow rapidly because of the crop-milk. Chicks fledge in 10 to 36 days (earlier if disturbed) and may continue to receive food from their parents for 30 to 40 days.
Parental Investment: altricial ; male parental care ; female parental care
Molecular Biology and Genetics
Statistics of barcoding coverage
Specimens with Sequences:852
Specimens with Barcodes:798
Species With Barcodes:121
Some species of pigeons and doves have expanded their ranges and increased their population sizes as a result of human activities (for example, 'rock doves Columba livia' and 'Eurasian collared doves Streptopelia decaocto'). Other species are less fortunate and their ranges and populations are shrinking as a result of habitat loss and fragmentation, hunting, introduced species, agriculture and pesticides. Columbids that live on islands are the most threatened. Habitat preservation is the best solution to dwindling numbers of some columbids, and captive breeding may be useful as a last resort.
The IUCN lists 109 species of columbids in various categories from ‘Extinct’ to ‘Near Threatened. CITES lists 26 members of Columbidae ranging from Appendix I to Appendix III.
Relevance to Humans and Ecosystems
Because they feed on cultivated grain, columbids are often thought of as crop pests. They are also pests in urban areas where they nest in man-made structures and their droppings can be a nuisance. They are also known to carry human disease.
Negative Impacts: injures humans (carries human disease); crop pest; household pest
Pigeons and doves are often a part of folklore and literature, and havae been domesticated for food (both eggs and adults are eaten by people). Research involving columbids has lead to increased knowledge about the inheritance of morphological and behavioral traits, endocrinology, learning, evolution, orientation and navigation. Pigeon racing is also a common pass-time and racing pigeons can sell for as much as $350 000. Pigeons and doves were also used as messengers during war times and are sometimes kept as pets.
Positive Impacts: pet trade ; food ; research and education
Pigeons and doves constitute the bird clade Columbidae, that includes about 310 species. Pigeons are stout-bodied birds with short necks, and have short, slender bills with fleshy ceres. They feed on seeds, fruits, and plants. This family occurs worldwide, but the greatest variety is in the Indomalaya and Australasia ecozones.
In general, the terms "dove" and "pigeon" are used somewhat interchangeably. Pigeon is a French word that derives from the Latin pipio, for a "peeping" chick, while dove is a Germanic word that refers to the bird's diving flight. In ornithological practice, "dove" tends to be used for smaller species and "pigeon" for larger ones, but this is in no way consistently applied, and historically, the common names for these birds involve a great deal of variation between the terms. The species most commonly referred to as "pigeon" is the feral rock pigeon, common in many cities.
Doves and pigeons build relatively flimsy nests – often using sticks and other debris – which may be placed in trees, on ledges, or on the ground, depending on species. They lay one or two eggs at a time, and both parents care for the young, which leave the nest after seven to twenty-eight days. Unlike most birds, both sexes of doves and pigeons produce "crop milk" to feed to their young, secreted by a sloughing of fluid-filled cells from the lining of the crop. Young doves and pigeons are called "squabs".
- 1 Taxonomy and systematics
- 2 Description
- 3 Distribution and habitat
- 4 Behaviour and ecology
- 5 Status and conservation
- 6 Relationship with humans
- 7 Respiratory and circulatory physiology
- 8 Relevant physical and chemical properties of pumping blood
- 9 Integration of respiratory and circulatory organs and effects on physiology
- 10 See also
- 11 References
- 12 Further reading
- 13 External links
Taxonomy and systematics
The Pteroclididae (sandgrouse) were formerly included in the order Columbiformes largely due to their reported ability to drink by the "sucking" or "pumping" action of peristalsis of the esophagus ("The only other group, however, which shows the same behaviour, the Pteroclididae, is placed near the doves just by this doubtlessly very old characteristic." ); more recently, it had been reported that they cannot drink by "sucking" or "pumping", and were treated separately in the order Pteroclidiformes and were considered to be closer to the shorebirds. Recent phylogenomic studies support the grouping of these pigeons and sangrouse together, along with mesites, forming the sister taxon to Mirandornithes.
Columbidae are usually divided into five subfamilies, probably inaccurately. For example the American ground and quail doves, which are usually placed in the Columbinae, seem to be two distinct subfamilies. The order presented here follows Baptista et al. (1997) with some updates.
The arrangement of genera and naming of subfamilies is in some cases provisional, because analysis of different DNA sequences yield results that differ, often radically, in the placement of certain (mainly Indo-Australian) genera. This ambiguity, probably caused by long branch attraction, seems to confirm the first pigeons evolved in the Australasian region, and that the "Treronidae" and allied forms (crowned and pheasant pigeons, for example) represent the earliest radiation of the group.
The dodo and Rodrigues solitaire are in all likelihood part of the Indo-Australian radiation that produced the three small subfamilies mentioned above, with the fruit-doves and pigeons (including the Nicobar pigeon). Therefore, they are here included as a subfamily Raphinae, pending better material evidence of their exact relationships.
Exacerbating these issues, columbids are not well represented in the fossil record. No truly primitive forms have been found to date. The genus Gerandia has been described from Early Miocene deposits of France, but while it was long believed to be a pigeon, it is more likely a sandgrouse. Fragmentary remains of a probably "ptilinopine" Early Miocene pigeon were found in the Bannockburn Formation of New Zealand and described as Rupephaps; "Columbina" prattae from roughly contemporary deposits of Florida is nowadays tentatively separated in Arenicolumba, but its distinction from Columbina/Scardafella and related genera needs to be more firmly established (e.g. by cladistic analysis). Apart from that, all other fossils belong to extant genera. For these, and for the considerable number of more recently extinct prehistoric species, see the respective genus accounts.
Pigeons and doves exhibit considerable variations in size. The largest species is the crowned pigeon of New Guinea, which is nearly turkey-sized, at a weight of 2–4 kg (4.4–8.8 lb) The smallest is the New World ground-dove of the genus Columbina, which is the same size as a house sparrow and weighs as little as 22 g. With a total length of more than 50 cm (19 in) and weight of almost 1 kg (2 lb), the largest arboreal species is the Marquesan imperial pigeon, while the dwarf fruit dove, which may measure as little as 13 cm (5.1 in), has a marginally smaller total length than any other species from this family. Smaller species tend to be known as doves, while larger species as pigeons, but no taxonomic basis distinguishes between the two.
Overall, the Columbidae tend to have short bills and legs, and small heads on large compact bodies. Their characteristic head bobbing was shown to be due to their natural desire to keep their vision constant in a 1978 experiment by B. J. Frost in which they were placed on treadmills – they did not bob their heads as their surroundings were constant. The wings are large and have low wing loadings; pigeons have strong wing muscles (wing muscles comprise 31–44% of their body weight) and are among the strongest fliers of all birds. They are also highly manoeuvrable in flight.
The plumage of the family is variable. Granivorous species tend to have dull plumage, with a few exceptions, whereas the frugivorous species have brightly coloured plumage. The Ptilinopus fruit doves are some of the brightest coloured pigeons, with the three endemic species of Fiji and the Indian Ocean Alectroenas being the brightest. Pigeons and doves may be sexually monochromatic or dichromatic. In addition to bright colours, pigeons may sport crests or other ornamentation.
Like some other birds, Columbidae have no gall bladders. Some medieval naturalists concluded they have no bile (gall), which in the medieval theory of the four humours explained the allegedly sweet disposition of doves. In fact, however, they do have gall (as Aristotle already realised), which is secreted directly into the gut.
Distribution and habitat
Pigeons and doves are distributed everywhere on Earth, except for the driest areas of the Sahara Desert, Antarctica and its surrounding islands and the high Arctic. They have colonised most of the world's oceanic islands, reaching eastern Polynesia and the Chatham Islands in the Pacific, Mauritius, the Seychelles and Réunion in the Indian Ocean, and the Azores in the Atlantic Ocean.
The family has adapted to most of the habitats available on the planet. These species may be arboreal, terrestrial or semiterrestrial. Various species also inhabit savannas, grasslands, deserts, temperate woodlands and forests, mangrove forests, and even the barren sands and gravels of atolls.
Some species have large natural ranges. The eared dove ranges across the entirety of South America from Colombia to Tierra Del Fuego, the Eurasian collared dove has a massive (if discontinuous) distribution from Britain across Europe, the Middle East, India, Pakistan and China, and the laughing dove across most of sub-Saharan Africa, as well as India, Pakistan and the Middle East. Other species have a tiny, restricted distribution; this is most common in island endemics. The whistling dove is endemic to the tiny Kadavu Island in Fiji, the Caroline ground-dove is restricted to two islands, Truk and Pohnpei in the Caroline Islands, and the Grenada dove is restricted to Grenada in the Caribbean. Some continental species also have tiny distributions; for example, the black-banded fruit dove is restricted to a small area of the Arnhem Land of Australia, the Somali pigeon is restricted to a tiny area of northern Somalia, and Moreno's ground dove is restricted to the area around Salta and Tucuman in northern Argentina.
The largest range of any species is that of the rock dove. This species had a large natural distribution from Britain and Ireland to northern Africa, across Europe, Arabia, Central Asia, India, the Himalayas and up into China and Mongolia. The range of the species increased dramatically upon domestication, as the species went feral in cities around the world. The species is currently resident across most of North America, and has established itself in cities and urban areas in South America, sub-Saharan Africa, Southeast Asia, Japan, Australia and New Zealand. The species is not the only pigeon to have increased its range due to the actions of man; several other species have become established outside of their natural range after escaping captivity, and other species have increased their natural ranges due to habitat changes caused by human activity.
Behaviour and ecology
Seeds and fruit form the major component of the diets of pigeons and doves. In fact, the family can be divided into the seed-eating or granivorous species (subfamily Columbinae) and the fruit- and mast-eating or frugivorous species (the other four subfamilies). The granivorous species typically feed on seed found on the ground, whereas the frugivorous species tend to feed in trees. There are morphological adaptations that can be used to distinguish between the two groups; granivores tend to have thick walls in their gizzards, whereas the frugivores tend to have thin walls. In addition, fruit-eating species have short intestines, whereas those that eat seeds have longer ones. Frugivores are capable of clinging to branches and even hang upside down to reach fruit.
In addition to fruit and seeds, a number of other food items are taken by many species. Some, particularly the ground-doves and quail-doves, take a large number of prey items such as insects and worms. One species, the atoll fruit dove is specialised in taking insect and reptile prey. Snails, moths and other insects are taken by white-crowned pigeons, orange fruit doves and ruddy ground doves.
Status and conservation
While many species of pigeons and doves have benefited from human activities and have increased their ranges, many other species have declined in numbers and some have become threatened or even succumbed to extinction. Among the 10 species to have become extinct since 1600 (the conventional date for estimating modern extinctions) are two of the most famous extinct species, the dodo and the passenger pigeon.
The passenger pigeon was exceptional for a number of reasons. It is the only pigeon species to have gone extinct in modern times that was not an island species. It was once the most numerous species of bird on Earth. Its former numbers are difficult to estimate, but one ornithologist, Alexander Wilson, estimated one flock he observed contained over two billion birds. The decline of the species was abrupt; in 1871, a breeding colony was estimated to contain over a hundred million birds, yet the last individual in the species was dead by 1914. Although habitat loss was a contributing factor, the species is thought to have been massively overhunted, being used as food for slaves and, later, the poor, in the United States throughout the 19th century.
The dodo, and its extinction, was more typical of the extinctions of pigeons in the past. Like many species that colonise remote islands with few predators, it lost much of its predator avoidance behaviour, along with its ability to fly. The arrival of people, along with a suite of other introduced species, such as rats, pigs, and cats, quickly spelled the end for this species and all the other island forms that have become extinct.
Around 59 species of pigeons and doves are threatened with extinction today, about 19% of all species. Most of these are tropical and live on islands. All of the species are threatened by introduced predators, habitat loss, and hunting, or a combination of these factors. In some cases, they may be extinct in the wild, as is the Socorro dove of Socorro Island, Mexico, which was driven to extinction by habitat loss and introduced feral cats. In some areas, a lack of knowledge means the true status of a species is unknown; the Negros fruit dove has not been seen since 1953, and may or may not be extinct, and the Polynesian ground dove is classified as critically endangered, as whether it survives or not on remote islands in the far west of the Pacific Ocean is unknown.
Various conservation techniques are employed to prevent these extinctions, including laws and regulations to control hunting pressure, the establishment of protected areas to prevent further habitat loss, the establishment of captive populations for reintroduction back into the wild (ex situ conservation), and the translocation of individuals to suitable habitats to create additional populations.
Relationship with humans
The pigeon has contributed to both World War I and II, notably by the Australian, French, German, American, and UK forces. Thirty-two pigeons have been decorated with the Dickin Medal for war contributions, including Commando, G.I. Joe, Paddy, and William of Orange.
Cher Ami, a homing pigeon in World War I, was awarded the Croix de Guerre Medal with a palm Oak Leaf Cluster for her service in Verdun and for delivering the message that saved the Lost Battalion of the 77th Infantry Division in the Battle of the Argonne, October 1918. When Cher Ami died, she was mounted and is part of the permanent exhibit at the National Museum of American History of the Smithsonian Institution.
A grand ceremony was held in Buckingham Palace to commemorate a platoon of pigeons that braved the battlefields of Normandy to deliver vital plans to Allied forces on the fringes of Germany. Three of the actual birds that received the medals are on show in the London Military Museum so that well-wishers can pay their respects.
The rock pigeon has been domesticated for hundreds of years. It has been bred into several varieties kept by hobbyists, of which the best known is the homing pigeon or racing homer. Other popular breeds are tumbling pigeons such as the Birmingham roller and fancy varieties that are bred for certain physical characteristics, such as large feathers on the feet or fan-shaped tails. Domesticated rock pigeons are also bred as carrier pigeons, used for thousands of years to carry brief written messages, and release doves used in ceremonies. White doves are also commonly used in magic acts.
In the Hebrew Bible, doves or young pigeons are acceptable burnt offerings for those who cannot afford a more expensive animal. In the Book of Genesis, Noah sent out a dove after the great flood to determine how far the floodwaters had receded. "Dove" is also a term of endearment in the Song of Songs and elsewhere. In Hebrew, Jonah (יוֹנָה) means dove. The "sign of Jonas" in Mat 16:1–4 is related to the "sign of the dove".
Jesus's parents sacrificed doves on his behalf after his circumcision (Luke 2:24). Later, the Holy Spirit descended upon Jesus at his baptism like a dove (Matthew 3:16), and subsequently the "peace dove" became a common Christian symbol of the Holy Spirit.
In Islam, doves and the pigeon family in general are respected and favoured because they are believed to have assisted the final prophet of Islam, Muhammad, in distracting his enemies outside the cave of Thaw'r in the great Hijra.
The Chaldean myth the Epic of Gilgamesh, Utnapishtim releases a dove and raven to find land; however, the dove merely circles and returns. Only then does Utnapishtim send forth the raven, which does not return, and Utnapishtim concludes the raven has found land.
Several species of pigeons and doves are used as food, and probably any could be; the powerful breast muscles characteristic of the family make excellent meat. Domesticated or hunted pigeon have been used as the source of food since Ancient Middle East, Ancient Rome and Medieval Europe. It is familiar meat within Jewish, Arab, Assamese cuisine and French cuisines. According to the Tanakh, doves are kosher, and they are the only birds that may be used for a korban. Other kosher birds may be eaten, but not brought as a korban. It is also known in Asian cuisines, such as Chinese and Indonesian.
In Europe, the wood pigeon is commonly shot as a game bird, while rock pigeons were originally domesticated as a food species, and many breeds were developed for their meat-bearing qualities. The extinction of the passenger pigeon in North America was at least partly due to shooting for use as food. Mrs Beeton's Book of Household Management contains recipes for roast pigeon and pigeon pie, a popular, inexpensive food in Victorian industrial Britain.
Respiratory and circulatory physiology
Organ and tissue organization
The respiratory system of pigeons is an avian system. This system lacks a diaphragm and instead compensates this by multiple air sacs in the body cavity. Air sacs in the respiratory system do not directly contribute to gas exchange. The primary goal of the avian respiratory system is to deliver oxygen to tissues and remove carbon dioxide.
Unidirectional flow of air to the lungs is facilitated by the air sacs. Because of the unidirectional flow, birds maintain a higher oxygen content in their lungs and have an increased metabolic rate.
Air flow depends on the pressure changes of the air sacs. A negative pressure causes air to enter the respiratory system, and a positive pressure causes air to leave the respiratory system. Microscopic tubules known as air capillaries are the site of gas exchange within the avian lungs.
The avian respiratory system includes cells known as "free avian respiratory macrophages" (FARMS) or "free respiratory macrophages" (FRM). FRMs are the first line of defence, protecting the blood gas barrier from pathogens that may cause potential damage. In a healthy avian respiratory system, FRMs are not present on the respiratory surface such as the air capillaries but are on the atria.
These macrophages are also present on epithelial tissue below the atria and septa. FARMs also assist with cleansing the air inhaled by the animal before reaching the air capillaries, which may consist of harmful particles.
There are many components that make up the circulatory system: The arteries carry blood away from the heart and to the body tissues. The arterioles direct blood to where blood is needed in the body. The veins carry blood back to the heart from the body tissues. They split into venules, smaller veins that have the same function as veins, and further into capillaries, which are responsible for exchange of nutrients, gases and waste products between the body and blood.
The major arteries include carotids, brachials, pectorals, pulmonary arteries, aorta, coeliac, renal arteries, femoral arteries, caudal and posterior mesenteric. Carotids transport blood to the head and the brain. Brachials carry blood to the wings. Pectorals carry blood to the pectoralis which are the flight muscles. Aorta takes the blood to the whole body, except for the lungs which are supplied by pulmonary arteries. Coeliac is a branch of the aorta that conducts blood to the organs and upper abdominal tissues. Renal arteries deliver blood to the kidneys and the renal system. Femoral arteries bring the blood to the legs. Caudal directs blood to the tail, and posterior mesenteric takes blood to the lower abdominal tissues and organs.
As to veins, jugular anastomosis lets the blood flow from right to the left side of the body while the head is turned to one side. Jugulars take blood away from the head and the neck. Brachials take blood away from the wings. Pectorals empty the pectoral muscles and the anterior thorax. Superior vena cava, also known as the precava, carries blood from the anterior parts of the body. Inferior vena cava carries blood away from the posterior parts of the body. The hepatic vein drains blood from the liver. The hepatic portal carries blood away from the digestive tract. Coccygeomesenteric and hepatic portal vein drain the posterior digestive tract. Femorals take blood away from the legs. Sciatic veins empty blood from the thighs. The renal and renal portal veins carry blood away from the kidneys.
Heart type and features
Columbia livia pigeons have a four chambered heart consisting of the right atrium, left atrium, right ventricle and the left ventricle. The pigeon heart is larger in proportion to the body size than the human heart is. The relatively larger size and faster heart rate enable the pigeon cardiovascular system to reach the high metabolic needs required for flying. The system works by transporting oxygen towards the tissues and taking carbon dioxide away from the tissues. Other functions include getting rid of metabolic wastes, this is important for maintenance of body temperature.
How the heart works
Blood is pumped into the heart from the body through the veins. The blood enters the right atrium, is transported through the right ventricle, and is then moved to the lungs where oxygenation of the blood takes place. The blood then travels to the left atrium, exits the heart and moves through the arteries to the body via the left ventricle.
Relevant physical and chemical properties of pumping blood
The oxygenated blood is pumped out to parts of the body by the left ventricle, where after giving up its life fuelling oxygen and collecting the carbon dioxide, it returns, as deoxygenated blood to the right atrium through the caval veins. Then the blood is shunted to the right ventricle which pumps it to the lungs though the pulmonary arch where the carbon dioxide is stored to be exhaled and a new load of oxygen is picked up. This newly reoxygenated blood goes back to the left atrium of the heart through the pulmonary veins. The blood is then shunted to the left ventricle and the cycle repeats.
Cardiac output can be estimated in birds from the Fick equation. The volume of oxygen taken in per unit time is proportional to the difference in oxygen content between arterial and venous blood. The degree of proportionality depends on the volume of blood pumped per unit time, or cardiac output. In the pigeon, activity creates an increase in demand for oxygen. Volume of oxygen intake is increased by increasing the cardiac output of the heart. An active pigeon increases cardiac output through an increased heart rate to meet the oxygen requirement. It can increase about six times; however, the stroke volume decreases in an active pigeon.
Integration of respiratory and circulatory organs and effects on physiology
Most of the physiological changes for both the respiratory and circulatory systems have to do with the energetic requirements needed for flight. Smaller birds heart rates double from the resting state when they are flying and in larger birds the heart rate can increase three to four times from the regular rate.
Flight muscles require a large amount of oxygen and in order to obtain the oxygen the blood must be kept moving rapidly around the system. This occurs when the blood flows into the atria at the end of its journey around the body, or to and from the lungs. The ventricles are the pumping power houses that send the blood off on to repeat the cycle.
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