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Why chickens? The success of the species The aim of the book Relating the practical to the theory CHAPTER 1 EVOLUTION & DOMESTICATION Junglefowl ancestors in Asia Domestication in China How fowl spread from China around the world
CHAPTER 2 ANATOMY & BIOLOGY Skeleton and muscles, adaptations for f light Respiratory system Digestive system Immune system Sensory system; vision, smell, taste, hearing, touch Feathers; feather structure, molting Comb and wattles; thermoregulation, coloration Reproductive system in male and female
CHAPTER 3 BEHAVIOR Behavior of embryos Feeding and drinking Social behavior; aggression, “peck orders,” f locking behavior, communication Rest and sleep Courtship and mating Nest building and parental behavior Feather maintenance Chickens behaving badly; feather pecking, cannibalisim, bullying
112 CHAPTER 4 INTELLIGENCE & LEARNING
Brain structures Early learning, imprinting Problem-solving abilities Learning from others
216 Appendix: Full Breed List Bibliography & Webography
134 CHAPTER 5 BREEDS OF CHICKEN
Common breeds and their general characteristics; ornamental, egg-laying, meat Fancy fowl and the chicken frenzy of the 19th century Breeding for combs, feathers, body shape, and other aspects of appearance Characteristics of current commercial strains Evolution of the Rock Cornish and the Leghorn The ornamental breeds The egg-laying breeds The meat breeds
I N TR O D U C TI O N
hickens are extraordinary animals, although often misunderstood and under-appreciated. They are the most successful avian species on our planet, with an estimated 16 billion chickens living in countries throughout the world. They have been valued, and sometimes even revered, for their fighting proficiency, their striking appearance, the companionship that they offer, and of course their role in providing meat and eggs for human consumption. They are far smarter than most people think. Indeed, observations of their complex social organization led us to coin the term “peck order” to describe features, not just of chicken, but of human, social behavior. Chickens were domesticated approaching 10,000 years ago in Southeast Asia, and were initially bred from the Common or Red Junglefowl (Gallus gallus). They were probably initially used for cockfighting and sacrificial or religious purposes. They spread into Europe over the next several thousand years, where the Romans bred
I N T R O DUC T I O N
them as food-producing livestock, thus beginning the first commercial poultry industry. By the 19th century chickens were widely distributed throughout the world, and they started to be intensively bred, not just for meat and eggs, but also to create new varieties with distinctive feathering, combs, and body shapes. Chickens have transitioned successfully from their origins as part of small f locks in the tropics to their current status as one of our major dietary sources of animal protein because they are so adaptable. They are omnivores, and so are able to consume a variety of food types, including berries, seeds, plants, and insects. Different breeds can tolerate cold climates; and others, extremely hot and dry climates. Chickens can be kept in f locks ranging from just a few birds to tens of thousands. They are relatively self-sufficient, mainly requiring some protection from predators. For all of these reasons, chickens play a special role, not just in commercial production, but also in small-scale farming in both developed and developing countries.
I N TR O D U C TI O N
A N ATO M Y & B I O LO G Y
What does the world smell, taste, feel, sound and look like to a chicken? To answer this question, it is important to understand the structure of the chicken’s unique sensory system. The chicken has excellent vision, well-developed senses of touch, vision, hearing, and possibly smell, but a comparatively poor sense of taste. The most important sensory organs are located on the head, but chickens also receive sensory input through their skin. The skin, and especially the skin on the body parts not covered with feathers like the comb, wattles, and feet, is well supplied with touch receptors. These are sensitive to movement, pressure, hot and cold temperatures, and other pleasant or painful stimuli. The beak The beak is the chicken’s most important tactile structure. With its hooked upper mandible, the beak is shaped for grasping and tearing insects and plants. The bones that support the upper mandible can slide forward and backwards, which means that chickens, unlike mammals, can open both their lower and upper jaw, allowing them to grasp very small items. The beaks of chickens may be tipped or trimmed to prevent them damaging one another due to aggressive behavior such as feather pecking or cannibalism, although this type of behavior is normally only a problem when a f lock is kept in confined conditions. The procedure requires conservative management as the removal of a beak portion risks painful damage to the chicken’s sensory ability, because it destroys many of the specialized touch receptors. This, along with the changed shape of the beak, can result in the chicken having difficulty grasping and manipulating some types of food, as well as removing parasites like ticks when grooming.
T HE COMB Combs vary in size and shape in different breeds. They help regulate body temperature and are important for social recognition (see p42–45).
T HE E Y E Chickens have excellent color vision (see box). Their eyes are large, so a sizeable image is projected onto the retina. This results in high visual acuity and good panoramic vision. Because of the structure and lateral placement of the eyes, chickens can view close objects on the ground at the same time as distant objects out to the side, without having to shift their focus.
T HE E A R Hearing is an important sensory ability, since chickens communicate by vocalizing, even prior to hatching. The external opening of the ear is not conspicuous, but it has a colored lobe and is surrounded by specialized feathers that do not obstruct sound transmission. The range of frequencies that chickens can hear is somewhat narrower than that of humans.
T E T R AC H ROM AT IC V I S IO N
Chickens are tetrachromats: they have four types of cones in their retina and see four primary colors–red, blue, green, and ultraviolet–rather than the three that we see. While the f lower below left appears to us as yellow with a black center, the chicken also sees blue-violet areas on the center, petals, and stem, as shown below right. Chickens have ultraviolet plumage markings, so ultraviolet vision is probably an important aspect of their social communication.
T HE TONGUE The tongue contains a bone and is adapted for collecting, manipulating, and swallowing food. Chickens have hundreds of taste buds on their tongue and oral cavity, compared to the tens of thousands humans have. The sense of taste seems mainly to help chickens reject potentially noxious foods rather than to choose preferred foods.
S EN S O R Y S Y S TEM
T H E N O S T R I L S ( N A R E S) While it was once thought that olfaction is not important to chickens, recent research shows that they have a good sense of smell and use it to discriminate between different foods, for example those containing varying levels of fat.
T HE BE A K The beak is the chicken’s primary tactile structure, used for manipulating objects, feeding, grooming, nest building, and in social interactions. It is bone covered with hard keratin, and is richly supplied with blood vessels, nerves, and touch receptors. In both the upper and lower beak these receptors are organized into a “bill-tip organ” to help the chicken make fine discriminations, for example between food and non-food items.
T H E WAT T L E S These fleshy structures are usually more prominent in males than females. Blood circulation through the wattles helps regulate body temperature (see p42–45).
F E M A L E R E PRODUCT I V E SYST E M
Modern hens are incredible egg layers; unlike their Jungle Fowl ancestors, which lay about 10–20 eggs a year, chickens lay hundreds of eggs, with hybrid commercial strains of chickens laying more than 320 eggs per year. Humans have changed the egg laying patterns of hens largely by genetically breeding them for a high rate of lay with very few pauses in laying between successive clutches of eggs. Because the photoperiod or day length strongly affects egg laying, the control of lighting also plays a role in stimulating egg production, as does providing the hen with a diet that is high enough in energy to support her increased nutritional needs. Regardless of their importance as a source of foods for humans, from an evolutionary point of view eggs have only one purpose, to protect and nourish the developing embryo. Structure The hen has only one ovary and oviduct, on the left side of her body. The oviduct has various regions, each with its own specialized function (see right). It terminates in the cloacal cavity and the muscular vent opening, through which the egg is expelled. The ovary grows rapidly once the hen reaches sexual maturity, eventually weighing about 60 grams. It contains thousands of small oocytes (female reproductive germ cells.) Once these begin to mature into follicles they grow rapidly, increasing 20-fold in size in just eight days, at which time they are ready to be ovulated. Only a single oocyte is triggered to begin growing per day, so the oocytes become mature sequentially rather than all at once—leading the ovary to look like a bunch of different-sized grapes. These “grapes” (ova) are egg yolks. They are the largest cells in the animal kingdom, and are rich in fats to serve as a source of nutrition for the developing embryo.
The hen’s single ovary and oviduct is placed on the left side of her body.
A N ATO M Y & B I O LO G Y
T H E PAT H OF E G G C R E AT IO N
M AGNUM R EGION
Ovum and albumin
ISTHUMUS R EGION
Ovum and shell membranes
UTERUS R EGION
VAGINA R EGION
Egg formation At the beginning of egg formation, the follicle ruptures to release the ovum into the abdominal cavity. At the same time, it secretes the hormones estrogen and progesterone, triggering pre-laying behavior (see p190–194). The ovum then begins its 24-hour journey down the reproductive tract. It first enters the ostium, and is moved by the action of cilia into the magnum region over the next half hour or so. The ovum remains in the magnum for about three hours, during which time the glands in the magnum coat it with egg white. This is composed of two proteins—albumin and mucin. The albumin creates a thinner layer of egg white, while the mucin creates a more viscous layer, the thick white. The egg white serves as a supply of water for the egg to prevent dehydration, and acts as a heat reservoir so that the embryo does not become chilled when the hen leaves the nest to feed and drink. Egg development The egg then moves to the isthmus region, where, during the next hour, the two clear, semi-permeable shell membranes are deposited. These inner and outer shell membranes, which contain the albumin to prevent it from leaking out of the egg, are made of the protein keratin, and separate at the large end of the egg to form an air cell. After the membranes are deposited the egg moves to the shell gland in the uterus, where the outer shell is added. The outer shell is constructed to provide mechanical protection to the embryo and prevent harmful organisms from entering the egg, while at the same time allowing gases and water vapor to permeate the egg. It is made of layers of crystallized calcium carbonate, protein, and a cuticle that helps prevent water loss from the egg. The egg’s shell has small pores that allow oxygen to enter, and wastes like carbon dioxide and water vapor to leave. Shell pigments are also deposited on the egg while it is in the uterus. The motion of the egg in the uterus and the amount of time that the egg remains in the uterus determine the coloration and patterning of the egg shell. Rapid movement results in streaked patterns of color, while very slow movement is associated with rounded spots of color. Egg laying The egg remains in the uterus for about 20 hours, at which time the sphincter between the shell gland and the vagina relaxes. The shell gland then contracts, and the egg is laid by passing through the vagina, cloaca, and vent. The egg can be held in the cloaca for several hours, although it is not often that long.It rotates there, to be expelled large end first.
R EPR O D U C TI V E S Y S TEM / FEM ALE
N E S T S I T E S E L E C T I O N
When a hen is preparing to lay her egg, she seeks a concealed nesting site protected from predators, other chickens, and the elements. She may investigate several sites before selecting one, which she then prepares by making a scrape in the substrate. If appropriate nestboxes are provided hens will usually lay in them, although they sometimes find other sites that they like better, which is a problem for both backyard and commercial poultry keepers. Exposing hens to nestboxes before they begin laying, and training them to use the boxes when they first come in to lay, can help minimize this problem.
B EH AVIO R
B EH AV I O R
Courtship and Mating
Chickens, like their Junglefowl ancestors, have a harem-polygynous mating system. This means that there is a dominant male that maintains a harem of females, and that this male is the one that sires most of the offspring in the f lock by preventing other males from mating. This system is probably not as rigid as that seen in Junglefowl—domestication and selection of chickens has resulted in them being more socially tolerant and promiscuous, especially when they are kept in large f locks with multiple males. Nevertheless, it is true that many features of their reproductive behavior still mirror those of their ancestors.
S ex u a l d e vel o p m e n t
Chickens must learn the characteristics of appropriate mates when they are young in order to grow into sexually normal adults. This process is known as “sexual imprinting” and occurs at around 10–12 weeks of age. That this knowledge is not innate is shown by the propensity hand-reared birds have to show sexual crouching or courtship in response to
humans rather than other birds. Breeding problems can result if roosters and hens are not reared together during this sensitive developmental period—roosters that encounter hens for the first time when they are older generally show less successful mating behavior, although this problem decreases as they gain experience. Cockerels reared separately from females may even have poorer semen quality. Hens, unlike many other species of birds, do not need to have males present in order to become sexually mature. It is this trait, of course, which has made chickens so useful for backyard and commercial egg production.
Ma t i n g B e h a v i o r i n H e n s
Although the elaborate courtship display shown by roosters ( fig. 1–4) might lead one to believe that hens are simply passive recipients of mating, nothing could be further from the truth. Given a choice, hens are highly selective in determining which roosters they will mate with. Hens encourage courtship by approaching or staying near desirable roosters, particularly socially dominant ones. If the hen does not know a particular rooster’s dominance status, she uses a variety of physical cues to assess his suitability. These include his eye color, spur length, comb color, and most importantly his
comb size. Since males infected with parasites have smaller combs, this may be a way for the female to assess not only the male’s dominance status but his health, and thus his reproductive fitness. Hens mate multiple times per day, although less frequently than roosters, who mate between 10 and 30 times a day. One reason for this less frequent mating in hens is that they have a specialized gland in their reproductive tract in which they can store sperm. These sperm can remain viable for several weeks. This sperm storage gland seems to operate on the “last-in-first-out” principle, meaning that the sperm from the last rooster that mates with the hen on a particular day is the sperm that will fertilize the most mature follicle in her ovary. The importance of this pattern, and the fact that the hen’s fertility is lowest when she lays her egg in morning, can be seen in the timing of mating, with most mating occurring in the late afternoon. This also means that the hen can actively solicit mating by the most desirable males late in the day, increasing the chances that they will fertilize her eggs. In fact, there is some evidence that a hen can even selectively expel the sperm of an undesirable male, like a low-ranking one, from her sperm gland. fig. 3
Ma t i n g B e h a v i o r i n Ma l es
Roosters mate many times per day, although individuals vary greatly in libido. They may mate more than once in succession with a particular hen, although their interest in that hen quickly wanes over time. This is called the “Coolidge effect,” based on an exchange that supposedly occurred between President
C O U R TS H I P A N D M ATI N G
RO O S T E R C OU RT S H I P R I T UA L
The courtship ritual can be elaborate. The rooster first performs a behavior called “tidbitting” to attract the hen (Fig. 1) by pecking the ground and giving a call that signals that he has found a desirable food—whether or not there is such food! The rooster then “waltzes” to the hen, dropping one wing to the ground and circling around her (Fig. 2). When the hen crouches (Fig. 3) he mounts her, grabbing the feathers at the back of her neck and treading on her back with his feet (Fig. 4). This causes her to evert her cloaca, allowing the rooster to mate. The whole ritual takes between one and four minutes. Most chicken breeds are diurnally sensitive, naturally only breeding in the Spring and Summer months.
M EAT B R EED S
12 lb (5.5 kg)
9.5 lb (4.3 kg)
REGION OF ORIGIN
Features Although named for the Brahmaputra River in India, little is known about this breed’s origins. It may have originated in India and be the same breed as the Chittagong, or it may be an American cross between the Chittagong and Shanghai. Brahmas were exported from America to Europe in the 1850s and became a standard breed shortly thereafter. They are characterized by their pea comb, small head, arched neck, broad body, fully feathered long yellow legs, short tail, and upright posture. They are hardy and adapt well to cold climates. Behavior Brahmas are considered docile, trainable, socially tolerant, and unaggressive birds. They do not fly, and also do not vocalize as much as other breeds. The females go broody. Use Primarily used for meat, although their slow growth means they are not suitable for commercial production. The hens are not as prolific as other breeds, laying only about 150 brown eggs per year, but they do continue laying during the winter. Brahmas are also popular show birds because of their intricate plumage. Breed conservation status Under observation
COLORS AND FEATHER PATTERNS INLCUDE
Black, Buff, Dark (Silver-Penciled), Light (Columbian), White. Red, Blue, and Partridge (above)
M EAT B R EED S
8.5 lb (3.9 kg)
6.5 lb (3 kg)
REGION OF ORIGIN
Features Developed in the 1870s, this breed is notable for its rounded body shape, full breast, broad skull, stout yellow legs, dense downy feathers, and large body size. There are now also popular bantam varieties. Ancestors probably include both Asian and Continental breeds like the Brahma, Hamburg, Chittagong, or Cochin. They were originally called American Seabrights, but the name was changed to Wyandotte, a corruption of the name of a Native American Indian tribe the Wendat, when the breed was admitted to the APA Standard. The Silver Laced (pictured) was the first variety but there are now many colors and feather patterns. Because they have a small rose comb and full feathering they are suitable for colder climates. Behavior Because of the large body size of standard Wyandottes, they are not strong fliers. They are considered a friendly and docile breed that adapts well to both confinement and range conditions. The hens tend to go broody and are good mothers.
COLORS AND FEATHER PATTERNS INCLUDE
Barred, Birchen, Black, Blackbreasted Red, Blue, Blue Red, Brown Red, Buff, Buff Columbian, Silver Laced
Use Meat and eggs; also popular as show birds. Hens mature quickly and lay 200–240 brown or tinted eggs per year. Breed conservation status Recovering
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