E’ in preparazione ATLANTE ANATOMICO FOTOGRAFICO della BECCACCIA ( Scolopax rusticula)
WORK in PROGRESS
A colour Atlas of ANATOMY of EURASIAN WOODCOCK (Scolopax rusticola)
ANATOMY OF BIRDS
first reference at
A COLOUR ATLAS OF AVIAN ANATOMY *****
AVIAN ANATOMY GLOSSARY
Provides an alphabetical list of technical terms used to describe the structure of the body.
and at :
^ : a warm-blooded animal with two wings, two feet, a horny beak and a body covered with feathers.
Spinal column: part of the nervous system with the spinal column.
Lung: saclike respiratory organ.
Kidney: blood-purifying organ.
Ureter: duct that carries urine from the kidney to outside the body of a bird.
Cecum: cul-de-sac of the intestine.
Rectum: last part of the intestine.
Intestine: last part of the digestive tract.
Gizzard: last pocket of the stomach of a bird.
Liver: bile-producing digestive gland.
Heart: blood-pumping organ.
Crop: pocket formed by the bulding of the esophagus.
Esophagus: first part of the digestive tract.
Trachea: first part of the respiratory system.
Buccal cavity: chamber of the mouth.
and for Physiology
Indications synthetic by general character
of AVIAN ANATOMY at :
Margaret A. Wissman, DVM, Dipl. ABVP
As an avian veterinarian, I often use words like choana and cloaca, and when I do, I can easily visualize in my mind exactly what these body parts look like, and it's hard for me to realize that often, the owner doesn't understand what I'm saying. Let's take a learning tour through the bird, both externally and internally, so that bird owners will have a better idea of the anatomy.
In some areas, birds are similar to other animals, and in others, they are absolutely unique. For example, the bird has several unique adaptations that enables it to fly, including feathers, air spaces in bones, a beak in place of teeth and lips and the bones of the hand fused to support flight.
The head of the bird contains several structures that may confuse bird owners. The bill or beak is known anatomically as the rostrum. Now bear with me as I use some technical terms to explain common anatomical parts, but I think it's important that you see the scientific terms for the bird's anatomy, even if you won't commit them to memory on sight! The horny sheaths of the upper and lower beak can be called the maxillary rhamphotheca (or rhinotheca) and mandibular rhamphotheca (or gnathotheca). If you tip the head back, you will see a fleshy area under the lower mandible, and this is called the interramal region or interramal space. The tongue and related structures are nestled in this region. Sometimes, the first time an owner sees this region, they panic and think a piece is missing out of the beak. But, be assured, this space is normal. The maxillary rhamphotheca carries the paired nostrils or nares. Inside the nares of most parrots is a small, round, brownish structure called the operculum. Next time you take your bird out, closely examine the inside of the nostril, and you may see the operculum, which is quite obvious in the cockatiel and Amazon parrot. This may be mistaken for a seed or dried hand-feeding formula in the nostril, but the operculum is supposed to be there. However, if a bird is suffering from malnutrition or chronic sinus infections, a rhinolith (a mass formed of desiccated secretions and debris) may cause a physical obstruction to proper breathing, and it may disfigure the nares. Rhinoliths must be differentiated from the normal operculum found inside the nostril. In some birds (owls, parrots and pigeons) there is a fleshy band at the top of the rhinotheca that contains the nostrils or nares. This is called the cere. The cere is swollen, highly sensitive and may be feathered. In the budgie, the color of the cere in a mature bird may indicate the sex of the bird.
Inside the bones of the bird head are the sinuses and concha. These are hollow spaces normally, but with infection, they may become clogged with liquid, mucus, abscess material or debris. One sinus is found behind the eye, which is why some birds with respiratory and sinus infection may develop swelling and discharge from the eye.
Inside of the mouth, the oropharynx, contains the tongue, glottis, choana, palate, salivary glands, esophagus, opening of the avian equivalent of the Eustachian tubes (the pharyngotympanic tubes) and larygeal mound. The tongue has a bone in it. The tongue is adapted for collecting food, manipulating food and swallowing. For example, the tongue of the birds in the lory and lorikeet families is the most specialized of the parrots. The lory tongue is called a '"brush-tongue," which refers to a cluster of elongated papillae that are normally only visible when the bird is feeding on liquid or soft foods, or when preening another bird.
At the base of the tongue, the glottis and the laryngeal mound are located. The larynx of mammals is used for vocalization, but it is the syrinx, located down much further, that is responsible for sound production in birds. The glottis is the opening to the windpipe, or trachea. The choana is located on the roof of the mouth. It is a slit that connects through some passages to the nostrils. One really neat difference that birds have is that the glottis will fit snugly into the choanal slit when the bird closes its mouth, and the bird will then have a closed connection from the nostrils to the windpipe. When a human breathes through the nostrils, the air goes through the back of the throat, which is an open area, to the trachea through the larynx. There are little projections, called papillae, that normally are found at the edges of the choanal slit. Other papillae, pointing towards the back of the throat, may be found in the oropharynx. A second, smaller slit is located behind the choanal slit. This is the opening to the middle ears, the infundibular cleft, of birds, connected by a tube, called the pharyngotympanic tube. Birds with middle ear infections often have a red, swollen cleft. This cleft is important for birds that fly at great altitudes, as it helps equalize pressure in the middle ear. I'll bet you never thought that birds ears might pop when they ascend during flight, like ours do when we humans fly!
In the past, it was thought that birds had a poorly developed sense of taste. Taste buds lie at the base of the tongue, in most species of birds. Some birds have taste buds on the inside of the tip of the lower and upper bills and there are several sites on the roof of the oropharynx, near the choana. In parrots, the taste buds are on the roof of the oropharynx on either side of the choana, and on the floor of the oropharynx at the front end of the laryngeal mound. Mallard ducks have less than 500 taste buds, compared to the 10,000 of a human and 17,000 in the rabbit. Birds do have a sense of taste, and do show definite taste preferences, as we all know!
Examination of the oropharynx is extremely important when a bird is evaluated by an avian vet. I use a strong, focal light source and magnification to closely evaluate the choana, papillae and infundibular cleft. It gives the experienced avian vet a tremendous amount of information regarding the overall health of the bird. If the choana is swollen, if the papillae are blunted or absent, if the infundibular cleft is reddened, if abscesses are present, or if thick, white ropy mucus is present, it gives the vet a good idea if malnutrition, vitamin A deficiency, bacterial or yeast infection or middle ear infections may be present, to name just a few. Internal papillomatous disease (papillomas) can occur throughout the gastrointestinal tract, and lesions may be present in the oropharynx. They may look like small, pink wart-like lesions. If your vet does not closely examine this area during a check-up, ask her why. It should be examined in every bird!
The oropharynx is variably colored in different species of birds. It may be pink, black or mottled. It takes experience to determine if the throat is blotchy pigmented, or if disease is present. For example, the oropharynx of the blue and gold macaw may be uniformly black, or it may contain pink areas. Pink areas could indicate internal papillomatous disease, or they could be normal. I always recommend that breeders get into the habit of examining the oropharynx of all of their baby birds daily, to look for changes that could be a problem. It is easy to begin learning the normals from the abnormals, if a breeder looks at the throats every day.
The rest of the head is divided into three regions, the forehead, crown and back of the head, and the three areas together are sometimes called the pileum. The orbital region is a narrow zone around the eye, and includes the eyelids. The ear region surrounds the ear opening, called the external acoustic meatus. The ear coverts are feathers that overlie the meatus, and have a different texture than the other head feathers. In most birds, the meatus is behind and slightly below the level of the eye. Next time you are scratching your bird's head, gently part the feathers in this area and take a peek at the external ear opening. There have been times when I was performing a new bird pre-purchase exam and I was the fourth or fifth vet to see the bird, and no one else had made a note of the fact that the bird was missing an external meatus on one side! This area is often overlooked when a bird is examined. Macaws that started out as stunted babies may have external acoustic meatus problems from infection or scarring. Lovebirds seem to be more prone to external ear infections (otitis externa) than other species of birds.
The meatus opening is often just a small hole, but in the owl it is huge. Some owls have a movable flap along the front edge, called the operculum.
Birds are almost all intensely visual animals. One indication of this is by the size of the eye, which is extremely large in relation to the rest of the head. Some owls and hawks have eyes that are absolutely as large or even larger than those of a human. The relatively large eye of the bird permits a correspondingly large image to be projected on the retina, which contributes to the excellent acuity of avian vision.
One interesting difference between the primate eye and the bird eye is that the primate lens filters out wavelengths of light below 400 nm which renders ultraviolet radiation invisible. However, the bird lens is optically clear and appears to transmit wavelengths of light down to about 350 nm, which makes near ultraviolet radiation visible to the bird, and absorbing only those ultraviolet wavelengths that are not dangerous to the bird. Current research indicates that birds may secrete a substance from the uropygeal (preen) gland that is spread on the feathers and is visible in the ultraviolet range. It is suspected that this may be one way that a bird may visually discern the sex of other birds, through the differences in ultraviolet color in the feathers. This has not been proven, yet, however. But it is known that birds can see the three visual pigments that normal humans can see, called trichromatic, and they may be tetrachromatic because they may be able to see in the ultraviolet range.
One would think that birds can see color, based on the spectacular plumage of many birds, such as the brightly colored lories, flashy peafowl and gaudy macaws. And this is correct. There are two different types of cells found in the retina of the eye, the rods and cones. Rods are sensitive to the intensity of light, so it makes sense that nocturnal (night active) birds such as owls have mostly rods. The cones are responsible for visual acuity the sharpness with which detail is perceived) and color vision. Diurnal (daytime active) birds have far more cones than rods.
Without going into detail, some birds have remarkable adaptations. For example, the eye of water birds and birds that live on open plains have a special area of the retina that can fix the horizon accurately as a reference point. Isn't that amazing? The anatomical characteristics of the diurnal avian eye make it likely that it could see an entire panorama as accurately as a mammal could see a single detail. For example, both a bird and a human should be able to see a mouse from a height of 250 feet, but the human could only do so if his attention were accurately directed to it, but the bird should be able to see it without looking directly at it. Also, the bird should be able to see all the mice in the field with a single glance, whereas the human could only do this by scanning the area laboriously. Birds can assimilate detail much faster than a mammal, and should be better able to detect and follow movement. Owls can gather light much more effectively than humans, and would be able to see a mouse at the opposite end of a football stadium in what we would perceive as pitch darkness. Diving birds that must see underwater have two separate areas on the retina for focusing, since the eyeball becomes deformed by the water pressure when they are underwater. Aren't birds just remarkable?
Birds have three eyelids, and so do dogs and cats. The upper and lower eyelids have small bristle feathers that resemble eyelashes. Most birds only close their eyelids during sleep, and use the third eyelid alone for blinking. The third eyelid, the nictitating membrane, lies beneath the eyelids on the side of eye closest to the nostril. It darts across the eye about 30-35 times per minute in the domestic fowl, and also moves across the eye if an object approaches the eye suddenly or if something touches the head. The third eyelid becomes scooplike and sweeps excess fluid in to the corner of the eye where it drains. In most birds, the nictitating membrane is transparent, so vision is not impaired when the eyelid blinks, which is important since so many birds are prey animals. It helps to be able to see when blinking! It is suspected that some birds may fly with the third eyelid covering the cornea of the eye, which prevents it from drying out during flight, acting like birdy goggles.
The iris is the colored portion of the eye, and the pupil is the hole that allows light in. The iris often changes color as a parrot matures at a predictable time. Many baby parrots have a grey iris, which is one easy way to determine age in a young bird. The iris of the African grey parrot and blue and gold macaw is grey when they are babies, and gradually changes to a yellow color when the bird is between 10 and 16 months of age, although this is variable. Baby Senegal parrots also have a grey iris, that changes to a yellow-orange color by about 12 months of age, however, it has been reported that the iris of parent-fledged Senegals raised outdoors in Florida will have an adult colored iris upon fledging.
The lore (from the Latin, lorum), is a narrow elongated area between the eye and the maxillary rhamphotheca. This area is directly in front of the eye, and in many species of birds (songbirds, native American birds) there is a straplike line, the loral stripe or loral line, beginning at the maxillary rhamphotheca and running to the eye area. Behind the eye area, this stripe continues as the eye stripe. These two stripes together form the lateral stripe. The lore not only carries a stripe, but it may have special feathers, such as loral bristles, or it may be naked. The Red-Lored Amazon, (not Red-Lord, as I often see it written in newspapers) Amazona autumnalis, and the Yellow-Lored Amazon, Amazona xantholora, are both named for the color found in the lore area on the head.
There are many different adaptations of the bill in birds that vary tremendously in structure depending on the functions necessary and the diet consumed. The bill functionally replaces the lips and teeth of mammals. Another unusual feature of the upper jaw of the bird is that it is moveable, unlike that of mammals. This is made possible by elastic zones in the bones of the face. Psittacine birds have a very strong beak and are also known as hookbills, for obvious reasons. Some parrots have rasplike ridges that run transversely inside the upper bill which can reduce the hardest pits to dust. If you peek up into the upper beak of a macaw, these ridges will be very obvious. Seed-cracking birds such as finches and canaries have a stout, conical beak. There are birds called crossbills, with sharply-pointed upper and lower components of the bill that cross over to hold the scales of fir cones apart so that the tongue can remove the seeds. Hummingbirds consume nectar and pollen and have a bill adapted to penetrate deeply into the throats of flowers. The Hawaiian goose has a specialized bill that can crop vegetation. Flamingoes and many ducks have a series of plates in the bill that filter out small organisms by straining water. Pelicans have an extensive interramal region that has developed into a pouch-like dip net. Anhingas can spear fish with a daggerlike beak. The beak of birds of prey is a hooked bill that is powerful and sharp-pointed. One very interesting bill belongs to the European Nightjar. It is a short wide bill with rictal bristles for netting insects while in flight. The rictal area is along the sides of the upper and lower mandibles, and in the Nightjar, they appear as a moustache that acts as a net to catch insects when the bird's mouth is open.
The bill resembles skin microscopically, because it contains dermis and epidermis. But the epidermis is very thick and contains calcium and keratin, which gives the bill its typical hardness. Although the bill of most birds is thick and hard, it is soft and leathery in some waders. The bill is only hard at the tip in ducks and geese.
The beak has a bill-tip organ in the upper and lower bills. This organ is very sensitive and is used by the bird to feel the environment, and allows the bird to discriminate between food and other particles. It makes sense that a bird like a duck or goose that must sift through the mud to find food will require a method of feeling with its beak like how we might use our sensitive fingertips. Because the beak is so sensitive, no one should EVER intentionally cut a beak back short enough to make it bleed as a method of "attitude adjustment." The bird relies on its beak like we use our fingers, so it is dangerous and cruel to cut a beak to the bleeding point. Overgrown beaks are like our fingernails. The dead portion of the fingernail and beak have no feeling, but may transmit heat and vibration. So, the overgrown beak may be safely trimmed back to normal length without causing pain to the bird. A good avian vet knows what a normal beak shape and length should be, so that it can be properly trimmed. The nerve endings for the beak are in channels that can be seen as white dots in a black beak.
The newly hatched baby bird has a small pointed hardened process on the front portion of the upper beak, called the egg tooth. The egg tooth is used by the hatchling to pip into the air cell of the egg, and then to break and unzip the shell. It is then shed sometime after hatching.
The neck is another interesting area of the bird. Some birds have very long necks (think of the graceful swan, the goose and the flamingo). Humans have seven cervical (neck) vertebrae, which are the bones that are surround the spinal cord. Birds have between 11 to 25 cervical vertebrae, which varies with the length of the neck. Nature designed the bird so that the minimum length of the neck is long enough to enable the bird to reach the uropygeal gland (also called the preen gland, found at the base of the tail in species that have one) so that it can properly preen. The neck is usually proportional to the length of the legs. So, long legs are generally found with a long necked bird, which enables the bill to reach the ground. But a long neck doesn't have to accompany long legs (think of the goose). The shortest necks are found in the small passerine birds. In the neck, the esophagus moves from the center of the mouth to the right side. This is the reason that some people recommend feeding baby birds from the left side of the beak, with the syringe pointing to the right, although, in reality, it shouldn't matter which side of the beak a baby bird is fed from. The trachea (windpipe) also curves to the right in the neck. Also found in the neck are the paired jugular veins, with the right jugular usually being much larger. Avian vets may use the right jugular vein to draw a blood sample.
Generally, the neck is carried in a double curve, which forms an "S". Since the forelimb has been completely committed to flight in birds (as a group, although some birds, such as the emu and ostrich cannot fly), the bill has assumed the ability to perform many functions normally carried out by the mammalian forelimb, such as grooming and nest building. Anyone that owns a parrot knows that the beak is used, along with the tongue, to explore its environment. A person unfamiliar with birds may pull away when a bird that has had a hand extended to it to be picked up, reaches beak-first towards the hand prior to stepping aboard. The novice may incorrectly assume that the bird is going to bite him, but the bird is just using the beak to test the waters, so to speak, prior to stepping up.
Since the neck forms an "S" curve, it protrudes forward in the front, above the level of the crop. Often, this may be mistaken for a tumor or abnormality in the neck, especially when the crop is empty and the bird is sitting comfortably. Because the neck has more vertebrae than a human's and mammal's, the avian neck is extremely flexible, mobile and strong. We've all seen how easily a owl can turn its head so much farther around than we can. When a bird is comfortably restrained by an avian vet, the head and/or neck is held. The neck is considered one of the strongest parts of a bird's body, and it is almost impossible to injure a bird by holding it by the neck (as long as the windpipe is not closed off), let alone break its neck, when it is properly restrained. Often, people think, when they pick up a limp, dead bird, that it must have broken its neck, because the neck is so limber. It rarely is the cause of death. Birds that fly into a window or other solid structure may die, often of a concussion or other trauma, but in all my years of practice, I have only seen two birds with fractures of the cervical vertebrae.
In the front part of the neck of parrots, the crop (or ingluvies) is found. It is actually an outpouching of the esophagus, the tube that carries food from the mouth to the stomach. Many people think that all birds have a crop, but some do not, including the gull and penguin. In the parrot, it is oriented transversely across the neck. In pigeons and doves, the lining of the crop is shed when they are feeding babies, for the first few days. This is called crop milk, and it resembles mammalian milk in that it is rich in fat and protein, however, it lacks carbohydrates and calcium, and contains no milk sugar (lactose). The crop, in baby parrots, is very large, and shrinks down as the bird weans.
The trunk is the whole body of the bird between the neck and the tail. It is divided into the thorax, abdomen and pelvis. The thorax is bounded by the rib cage, sternum (keel) and vertebral column (backbones). The abdomen and pelvis aren't separated by any well-defined boundaries. The top part of the trunk is divided into the back and rump. The region between the right and left shoulder blades (scapulae) is called the interscapular region, and often carries distinctive streaks or colors. The whole back, combined with the top surface of the wings, is called the mantle. Often these anatomical descriptions are used by judges during bird shows. The side area of the trunk is called the flank. The underside is divided into the breast, belly and undertail. Another area, the crissum, refers to the general area around the vent, along with the undertail covert feathers. The term, vent, should only be used to describe the actual orifice, and not the general area under the tail.
The tail contains the flight feathers called retrices (which is Latin for rudders). The retrices are always paired, with the central one lying on the midline. The majority of birds have six pairs of retrices, but the number ranges from 6 to 32. Tail coverts are small feathers that lie over and under the retrices. Interestingly, the coverts are greatly enlarged in the peacock, and form the eyed feathers of the train. The pygostyle is the end-most bone of the spinal column. You may be more familiar with the term "the Pope's nose" for this extraneous piece found on a chicken thigh. If this bone has been fractured or injured, a male bird may not be able to successfully copulate with the hen.
Moving on to the wing, we find that it is unique, as it is adapted for flight. Many of the bones have become fused, and the skeleton of the hand (manus) has undergone considerable simplification. In addition to the bones of the wrist being consolidated, there are only three "fingers." The smallest one is called the alula, and the other two are called the major digit and minor digit. These "fingers" have reduced phalanges (our fingers have three phalanges each, except the thumb, which has two). Most commonly, the alula has one phalanx, the major digit has two, and the minor digit has one. The propatagium is that elastic triangular fold of skin on the leading edge of the wing. This is the area where a tattoo is placed when a bird is surgically sexed.
The feathers on the wing are divided into flight feathers (remiges), further divided into primary and secondary flights and coverts. The primaries are the last ten wing feathers on the wing, and are numbered from ten to one, outermost to innermost. The secondaries begin at the first bend of the wing, and are numbered one to twelve, from the bend inward. When wings are clipped, it should never be necessary to cut any feathers other than primaries, no matter which method is used.
The bones of the leg have also been modified. The thigh bone (femur) is the same as is found in mammals. The knee joint follows at the joint below the femur. The next bone is different from that found in mammals. Several bones have fused to form the tibiotarsus bone. Basically, the ankle bones fused with the bones of the arch of the foot to form one long bone. There is still a small fibula bone present. The next joint is called the intertarsal joint, and humans don't have one. The bone below that joint is called the tarsometatarsus, which also consists of fused bones. Think of the flamingo leg and how when you watch one walk, it seems as if the "knee" is bending the "wrong" way. That's because the elongated tarsometatarsus looks like the shin bone, so you think that the joint (if it was the knee) should bend in the other direction. But actually the knee is up in the feathered area of the leg, which DOES bend the same way ours does.
In most birds, the digits (toes) I through VI are present (we have five digits, numbered I through V). In most birds, the first toe is usually directed backwards, and the other three point forward, and this is technically known as the anisodactyl foot. However, in parrots, digits II and III point forward, and digits I and IV are directed backwards. This type of foot is called the zygodactyl foot. Swifts have a foot adapted for climbing, and all four toes point forward, and this is called the pamprodactyl foot. Emus, rheas, many wading birds and some woodpeckers only have three functional toes, and in most of these birds, it is the first digit (the hallux, which is similar to the big toe of man) that is lost. The ostrich has only two toes, with digits I and II being lost.
Some of the bones of the avian skeleton are hollow and connected to the air sacs of the respiratory system. Most of the vertebrae, pelvis, sternum and rib bones are hollow and the marrow has been eliminated. The limbs vary in the degree of pneumaticity, and there are pneumatic spaces within the bones of the head, as well. This is important to reduce the weight of a bird to allow it to be light enough for flight.
The skin of birds is different from other animals in several ways. For one, only birds have feathers. Some birds have ornamental outgrowths, characterized by thickened skin that has many blood vessels. For example, some birds have a comb, a bright red vertical projection from the forehead and crown. Some birds also have wattles, which are naked folds of skin that hang down from the mandibles. Some birds have ear lobes, which are folds of skin, that may be red, white or purple. The snood, also called the frontal process, and is a distensible fleshy process arising on the head between the eyes and nostrils of the turkey. Turkeys also have caruncles, which are small protuberances of skin on the head and upper neck.
The skin of birds contains no sweat glands, so birds rely on evaporative cooling from the respiratory tract. The main gland of birds is the uropygeal gland, and is present in most birds and may be relatively large in some aquatic species. It is absent in the emu, ostrich, many pigeons, Amazon parrots, and the hyacinth macaw, for example. Another adaptation with avian skin is the brood patch, an area over the breast that becomes thickened, very vascular and the feathers are lost during the brooding period. These modifications promote the transfer of heat from the hen to her eggs.
There are seven types of feathers, the contour, semiplume, down, powder down, hypopenna, filoplume and bristle feathers. The contours cover the surface of the body, and arise from feather follicles. The follicle consists of a living part and a nonliving part. Growing feathers, or blood feathers, have an active blood supply until the feather is grown out completely. If a bird plucks out a feather from a follicle repeatedly, it my eventually destroy the living portion of the follicle, resulting in a follicle that can no longer grow a feather. Feathers molt out when a new feather is developing in the follicle and the old feather is then pushed out. Normally, a plucked feather will begin to regrow from the follicle immediately, but a cut feather will not be lost until it is molted out. Most birds replace all their feathers yearly, and this is a continuous process, however, some birds molt during a particular time frame (for example, after breeding season, or in the summer).
The skin of birds has distinct, well-defined tracts called pterylae that contain the feather follicles for contours. The bare spaces between the pterylae are called apteria.
The digestive system has some unique avian features. We have already talked about the crop, the outpouching of the esophagus. The esophagus connects to the crop and then travels through the bones at the top of the keel. The esophagus then connects to the stomach. The avian stomach is unique. The first portion of it is called the proventriculus, and this is the part with glands in it that secretes gastric juice. The second part of the stomach is called the ventriculus, or gizzard, and it is where digested proteins are broken down and where grinding occurs. When a bird has PDD, proventricular dilation disease, the nerves to the gastrointestinal tract are usually affected, and the proventriculus will become dilated, thin-walled, and impacted with food items. The ventriculus may also become more mushy and less muscular.
Some birds have paired ceca (our appendix is really called the cecum, which is the singular of ceca). The big difference occurs when we examine the tail end of the digestive tract. Most people know that birds only have ONE external opening, called the vent, and the internal chamber, or cloaca, that is used for urination, defecation and reproduction. Inside the cloaca, there are three separate compartments, called the coprodeum, the deepest compartment, and is the terminal end of the rectum. The next cloacal compartment is the urodeum, and this is the middle section that collects urine and urates from the ureters, that drain the kidneys. In the hen, the left oviduct opens into the urodeum. When an egg is travelling through the reproductive tract of the hen, it enters the urodeum before it passes out through the vent. The last compartment, just inside the vent, is the proctodeum. This compartment contains the avian phallus, if one is present. The phallus differs from the mammalian penis in several ways, the major ones being that it is just a copulatory organ and that is does not function to drain urine. Male ratites (ostrich, emu, rhea), ducks, geese, swans and some domestic fowl and turkey possess a phallus. Male parrots do not. The proctodeum is also the site of the Bursa of Fabricius, an organ that produces cells to fight infections. The bursa is unique to birds and should always be harvested for histopathology if a baby bird dies, because it is often very helpful in diagnosing many avian conditions and diseases. The bursa begins regressing, called involution, several months after hatching, and will usually be completely involuted by sexual maturity, in most birds.
The urinary system of birds is different from mammals, as birds produce both urine and urates. The kidneys possess two different types of nephrons, the units that filter the blood to remove toxins and products of metabolism. Birds cannot concentrate their urine as well as mammals can. Birds also are uricotelic, meaning that they excrete the end product of nitrogen metabolism as uric acid, which is made in the liver and they excreted from the blood. Uric acid is the creamy white portion of the dropping. Urine is the clear portion. The feces constitute the third portion of a dropping, and this consists of the solid portion, usually brown or green, depending on what the bird has been eating. A bird is able to urinate independently of defecating, or passing feces, but most of the time, the bird will pass urine, urates and feces at the same time. And now we know the compartments where these are stored prior to being passed, right?
The respiratory system is very unique in birds. Although birds possess a larynx, as we do, they do not use theirs for producing sound. The syrinx is an organ found at the junction of the end of the trachea (windpipe) with the beginning of the large left and right primary bronchi. These are air tubes that allow the passage of air into the deeper portions of the respiratory tract. The lungs are found inside the bony ribcage, but not where they are located in a mammal. In mammals, they are found on either side of the heart, and have lobes. No bird lung has lobes, and the liver lies on each side of the heart, instead of the lungs. The liver is large in birds and is composed of a right and left lobe. Most parrots don't have a gallbladder. Bird lungs are located more against the bones of the back, and are relatively non-expansible. They are smaller by about 25% than those found in mammals. Birds do not have a diaphragm, as mammals do. Birds breathe by expanding the ribcage outward, which draws air in like a bellows. For this reason, it is vital that the chest not be prevented from moving outward when a bird is being restrained for examination or wing clipping, or it will not be able to breathe.
Without going into the detailed anatomy and physiology of the avian respiratory system, I want to mention a few differences between birds and mammals. Birds have paleopulmo and neopulmo, which are two systems of connections between the air tubes, the bronchi and parabronchi. The neoopulmo system is absent in primitive birds, such as the emu and penguin. In the place of alveoli, which are present in the mammalian lung, birds possess air capillaries instead. The air capillaries are closely entwined with a profuse network of blood capillaries. This is where gasses are exchanged. Birds also have an air sac system. The air sac is connected through a hole in the lung called an ostium. Some bone cavities are occupied by outpouchings of the air sacs.
There are eight air sacs in most species of birds. There are one cervical and one clavicular air sac, and two cranial thoracic, two caudal thoracic, and two abdominal air sacs. Occasionally, an air sac may rupture, and the bird may develop air under the skin (subcutaneous emphysema) or a large swelling of air in the neck region. When a bird is surgically sexed, the left caudal thoracic air sac is entered by the endoscope. The air sacs allow for easily visualization of the internal organs, usually, and the membrane between the caudal thoracic and abdominal air sacs may need to be punctured by the scope, to better visualize the gonad. Usually the scope in entered to be able to visualize the left gonad. If the trachea is blocked in a bird having breathing difficulties, or if surgery must be performed around the head and neck, it is possible to insert an air sac tube into one of the air sacs, allowing the bird to breathe through that, instead of the trachea. Our breathing only takes one breath to completely exchange the air in our lungs, while it takes a bird two breaths to completely exchange the air in the system. This is why an air sac tube can be used for breathing in a bird.
The female reproductive system is unique in birds. In most birds, the hen only has a left ovary and oviduct. However, two fully developed ovaries are usually present in birds of prey and the kiwi. Two oviducts may occur in birds of prey. In normal parrots and softbills, there is usually only a left ovary and oviduct. The ovary contains all of the cells that can turn into eggs. When ovulation occurs, the egg cell and yolk are released from the ovary. Since hens produce eggs, and do not develop the babies inside the uterus, as mammals do, the oviduct is the organ that receives the egg, and then applies the egg white, membranes and shell. There are five portions to the oviduct and each performs a different function. They are the infundibulum, which receives the egg after ovulation. This is where fertilization usually occurs. Sometimes a hen will lay eggs without a male being present, and in this case, the eggs will be infertile. Chicken eggs purchased at the grocery store are infertile eggs.
The next portion of the oviduct is the magnum. It is in this area that the bulk of the egg-white protein is added. The egg travels next into the isthmus where egg membranes are produced and calcification of shell begins. The next portion of the oviduct is called the uterus, but it is nothing like the uterus of a mammal. The uterus is also called the shell gland, as this is where the shell is put on the egg. The final portion of the oviduct is called the vagina, and it is here where the sperm are stored in the hen once copulation has occurred. It takes about 25 hours for the egg to travel down the oviduct. The oviduct terminates in the urodeum.
Unlike the hen's reproductive tract, the male usually has two functional testicles. However, they are located up inside the body near the kidneys, and are not found externally as they are in mammals. This is why most birds cannot be sexed by looking at the external characteristics of it, because the testicles or ovary are inside. Some birds have characteristics that identify it as male or female, such as different coloration of feathers. Eclectus parrots are an extreme example of this, as hens are predominantly red and purple and males are predominantly green. Other birds have more subtle differences. Birds that can be visually sexed are called sexually dimorphic. Monomorphic birds cannot be sexed by sight and must be sexed by chromosome analysis, DNA analysis or by endoscopy.
People often ask me about the heart in birds. It is relatively much larger than that of mammals. It also beats much faster than a mammal's. The heart of a bird can pump much more blood than a man or dog (about seven times as much!) The blood pressure is much higher in a bird, and it has a remarkable exercise capacity. The heart has four chambers, just like we do.
Blood cells in birds are different from those found in mammals. In birds, the red blood cells still contain a nucleus, and are very large in size, with a lifespan of only 20-35 days (120 days in man). Humans and other mammals have a type of white blood cell called the neutrophil, but in birds, these are called heterophils.
True lymph nodes, which are common in mammals, are not found in birds, except for certain aquatic birds. Birds and mammals do both possess a thymus, which is another organ of the immune system.
Birds possess remarkable abilities to navigate. Pigeons are notorious for their ability to find their way home when taken to strange, far-off locations and released. It is certain that the sun and stars are dominant orientational cues for birds. These provide compass information only. There is evidence that birds can tell the time of day by using their circadian rhythm with the sun's azimuth (direction from the observer). However, this does not explain how homing pigeons can orientate themselves accurately towards home from release points that are far distant and unfamiliar even when there is complete cloud cover. It is interesting to note that pigeons will miss their home base if they have been shifted six hours out of phase with true sun time. Pigeons may use polarized light or ultraviolet light to help them navigate, and they may use their sense of smell, as well. Homing pigeons can also detect sound frequencies below 1 Hz, which is much lower than many animals can detect. Birds may be able to detect the infrasound of waves breaking on the shore or wind whistling through mountain tops while flying, helping in navigation. Birds are also extremely sensitive to magnetic changes, and magnetic material has been found in a small area in the head of pigeons, which can aid in a bird locating the direction of the earth's magnetic field. Gravity and barometric pressure may also be used as clues to aid a bird in navigation. I am convinced that birds are very sensitive to environmental changes. For example, a friend has a pet Quaker parrot, Gaspar, and she had recently moved the cage to a location directly under her television set. The bird began feather picking immediately, and acted very agitated while the cage was in this location. Now, I'm no electrical expert, but I know that certain waves emanate from a television set, and this apparently really bothered Gaspar. Once she identified the cage location as a potential problem, she moved it, and voila, the picking ceased immediately. There are many mysterious features about birds that we have yet to learn, that's for sure.
Birds are fascinating creatures and possess many unique anatomical characteristics. I hope you now have a better understanding about the anatomy of these beautiful animals that we share our lives with.
by Linda Pesek DVM, Diplomat ABVP (Avian)
Digestion in birds involves a lot of organs, each performing a specific function. It begins with entry of food via the beak and ends with the exiting of refuse at the vent. Food is broken down and absorbed for use along the way.
The avian beak or rostrum is composed of bone and an outer horney Keratin sheath or rhamphoteca. Keratin replacement continues throughout the life of the bird. In large parrots, the keration is worn down by digging, eating and chewing on hard objects and is completely replaced in 6 months on the upper bill and in 2 - 3 months on the lower bill. The upper beak is the maxillary rostrum and the lower beak is mandibular rostrum. The cutting edges of the beak are tomia, which take the place of teeth. In some parrots, the underside of the upper beak has ridges for holding food and maintaining the edge of the lower beak. The tip of the upper and lower beak is richly supplied with nerve endings.
Beaks replace the lips and teeth of mammals and vary in shape, size, length and function according to the type of diet consumed. Seed-crackers such as finches have a short conical beak, while birds of prey such as hawks have a powerful hooked beak for tearing flesh.
The choana is a median fissure in the palate which connects the oropharynx inside the mouth with the nasal cavity. Birds do not have a soft palate as mammals do. Numerous projections or papillae are found at the edge of the choana. These papillae often become blunted with a vitamin A deficiency.
A small slit like opening, the infundibular cleft is located behind the choana. This permits communication between the pharanyx and eustachean tubes of the ears. Abundant lymphoid tissue is found in the wall of this infundibular cleft.
The tongue, just as the beak, is adapted to the type of food the bird consumes. Woodpeckers have a long narrow protrudible tongue which functions as a spear - allowing them to extract insects. Parrots, birds of prey, and finches have short, thick, fleshy tongues which allow them to manipulate their food. Fowl and pellicans have nonprotrusible tongues with caudally-directed papillae which allow the food to be easily shoved to the back of the mouth for swallowing. Birds do not have an epiglottis to prevent food from entering their larynx. Instead, the opening of the glottis closes during swallowing.
Numerous small salivary glands are found in the oral cavity. These are best developed in birds that eat a dry diet such as seed and insect eaters, and least developed in birds that eat a moist diet such as fish. Mucus is the major substance secreted by the avian salivary glands. It acts as a lubricant in swallowing.
The epithelial lining of the mouth in some species - especially passerines (songbirds) - is brightly colored - - which stimulates the parents to feed the chick when it begs for food.
The esophagus transports food from the mouth to the stomach. Many species of birds - such as parrots have an enlarged area of the esophagus known as a crop or ingluvies. Several types of crops exist. Gulls and penguins do not have a crop, while ducks, geese and song birds possess a small fustorm dilitation of their esophagus. The crop stores food temporarily and allows food to be softened before it enters the stomach. Pigeons and doves produce "crop milk" that they feed to their young for the first two weeks after hatching. Other species - such as parrots - will regurgitate food that has been stored and softened in their crops to their young.
Birds have a two part stomach, a glandular portion known as the proventriculus and a muscular portion known as the ventriculus or gizzard. Hydrochloric acid, mucus and a digestive enzyme, pepsin, are secreted by specialized cells in the proventriculus where chemical digestion begins. The gizzard has a thick muscular wall and plays an important part in the mechanical digestion by crushing and grinding food. The gizzard is lined by a cuticle or hardened carbohydrate and protein complex.
Fish and meateaters have a thin-walled sacklike stomach that is adapted for storage rather than mechanical digestion. The distinction between the proventiculus and the ventriculus is difficult to determine. Seedeaters (such as parrots), insectivores and herbivores have a very well developed gizzard adapted for mechanical breakdown of indigestible food. The proventiculus is easy to distinquish from the ventriculus in these birds.
The small intestine is the primary site of chemical digestion and nutrient absorption. It is divided into a duodenum, jejunum and ileum. These are arranged in u-shaped loops in the abdominal cavity. The large intestine is composed of paired caeca and a short rectum. The shape and size of the caeca vary in different species of birds. The caeca of the chickens are well developed while those of the passerines (song birds) are very small. Caeca are very rudimentary or absent in some species such as parrots and pigeons. Caeca contain lymphoid tissue. Cellulose breakdown by bacteria occurs in the caeca.
Chemical digestion and absorption of food occurs in the small intestine. Water resorption and temporary storage of waste material occurs in the cloaca and rectum.
The pancreas lies in the duodenal loop. The pancrease produces amylase, lipase and proteinases including trypsine. These are important enzymes for chemical digestion. It also produces the hormones insulin and glucogon which regulate blood sugar.
The large liver is divided into a right and left lobe. In most species the rught lobe is larger than the left. In some species - such as passerines - the right lobe is subdivided. The liver has numerous functions among which is the production of bile. Bile is important in the breakdown of fat. Bile is stored in the gallbladder.
Some species, such as parrots and pigeons, lack a gallbladder. In these species, bile is releaed directly into the small intestine.
The digestive tract is sterile at hatching. Altricial chicks - or chicks born blind and helpless - are innoculated with micrflora when parents feed them. Precocial chicks - chicks that are able to feed on their own shorly after hatching, are innoculated by bacteria from ther environment during feeding. In addition, a process known as "cloacal drinking" occurs in altricial birds by which microflora are "sucked" into the vent from the nest to colonize the posterior digestive tract.
Winged Wisdom Note: Dr. Linda Pesek graduated from the University of Pennsylvania School of Veterinary Medicine. She has a small animal and avian practice in New York. Linda also writes columns for The Long Island Parrot Society and The Big Apple Bird Club and is a frequent lecturer at their meetings. She is the owner of an extensive collection of exotic birds.
by Linda Pesek DVM
One of nature's true marvels is the design of the avian eye. In general, birds possess some of the best vision capabilities in the animal kingdom. Birds see in color which is important in recognizing food and danger and is possibly used in mating rituals. Birds also see with enormous accuracy and at long distances. Some have very advanced depth perception and motion detection capabilities, while others see well at night. A bird's eyes are quite large in comparison to the size of its body and are extremely important to its survival.
Nocturnal birds such as owls make very efficient use of light. Other birds, have finely tuned focusing ability, enabling them to see accurately at long distances. In fact some birds of prey (such as raptors) have developed enhancements to the eye which enable them to focus at both far and near distances and to accurately judge the speed of moving objects.
The following examples show various ways that humans have attempted to use this superior ability.
After World War II, the military experimented with using birds to help locate flyers downed in the sea. The birds could see downed pilots at much greater distances than did the human observers.
Companies that make pills (like asperin) or some candies, need final inspectors on the assembly line to remove the broken ones and to leave only perfect ones for packaging for the consumer. Human workers doing this type of work need frequent and long rest periods. A test using birds for this task proved that birds could work longer and be much more accurate.
Geese have been taught to eat the bad insects from crops while leaving the good insects alone. Their eye sight seems to be suited to this task.
The position of a bird's eyes on its head varies by type of bird. Birds of prey have wider heads with the eyes set apart and facing forward, similar to humans. This gives them binocular vision or depth perception and enables them to judge the speed and distances of prey and other objects.
Birds such as parrots, which are prey for other animals, usually have eyes set on the side of the head. This gives the bird better peripheral vision and enables it to detect danger from a wider field of vision. Birds can move their eyes a bit, to extend their range of vision, but do not have the extensive ability of some reptiles to rotate the eyes.
There are a number of diseases which can affect the eye, such as cataracts, conjuntivitis and absesses. Because birds rely so heavily on their eyesight, any problems should be treated immediately.
Although the anatomy of the avian eye is similar to that of mammals, a few reptilian characteristics remain. These reptilian characteristics are the pectin structure within the globe of the eye and the scleratic ring of bony plates - scleral ossicles that support the shape of the globe. Birds have enormous eyes relative to their body size and the keenest vision of all vertebrates.
The shape of avian eyes may be flat, globose, or tubular and varies according to the type of bird. "Flat" eyes are found in diurnal (awake during the day) birds with narrow heads such as doves, pigeons and parrots. They have a short anterior-posterior (front to back) axis, so that the image that falls on the retina is small, resulting in lower visual acuity. "Globose" (more rounded) eyes are found in birds with wider heads such as raptors and passerines (canaries, finches). The anterior-posterior axis is greater than in flat eyes, increasing visual acuity (focusing quality). "Tubular" eyes are characteristic of nocturnal birds of prey, such as owls.
The postion of the avian eye varies according to the shape of the skull. Pigeons, doves and parrots have narrow skulls with laterally placed eyes. Birds of prey, such as raptors, have broader skulls with more frontally positioned eyes. Laterally positioned eyes provide a large field of vision but a small binocular vision. Binocular visions means that both eyes are focused on the same object and move in a coordinated fashion.
The eye is composed of three "coats" or layers. The outer coat is the sclera. This is a strong white fibrous coat which maintains the shape of the eye, protects its internal structure, and serves as the attachment point for the eye muscles. The clear front portion of the eye is the cornea. Light passes through the cornea and into the pupil behind it. A series of small, overlapping bones - the scleral ossicles - are located in the sclera encircling the cornea. These bones strengthen the eyes and provide attachment for the ciliary muscles.
The middle coat or layer of the eye is the vascular tunic. It is composed of the iris, the ciliary body and the choroid. The iris is a thin sheet of muscle fibers and connective tissue that forms a diaphragm in front of the lens, controlling the amount of light entering the eye. The pupil is the circular open space at the center of the iris. The iris contains pigment that gives the eye its color. In some species of birds, the iris coloration changes with age or sex of the bird. For example, white and pink female cockatoos have a red iris, while males have a dark brown to black iris.
The ciliary body suspends the lens by zonular fibers. The ciliary body is located at the base of the iris. It suspends the lens and alters the shape of the lens, changing the focal point of the eye. Ciliary processes or folds formed by the ciliary body produce a thin fluid known as aqueous humor. This fluid is located between the cornea and the iris and between the iris and the lens. It maintains intraocular pressure and helps maintain the global shape of the eye. The ciliary muscles are very important for visual accomodation. The choroid is a thick, vascular, darkly pigmented layer that coats the retina and helps nourish both the sclera and retina.
The third coat or innermost layer of the eye is the retina. It is very sensitive to light and contains photo receptive rods and cones. Cones are important for visual acuity and color vision and are most numerous in nocturnal birds. The retina of birds is different from mammals in that it is thick and has no blood vessels. The pecten is a vascular, folded structure projecting from the avian retina. It is thought to aid the choroid layer in supplying the retina with nutrients and in transporting oxygen and carbon dioxide. The optic nerve centers the retina in the lower quadrant of the back of the eye.
The area of sharpest visual accuity of the retina is in the macular disk. The fovea centralis is a tiny pit in the center of the mocular disk. It is the area of maximal optical resolution. Most birds have single fovea. Birds that pursue fast moving prey - such as falcons - possess two foveate areas in each eye giving them very accurate perception of distance and speed.
The avian lens is softer than that of mammals - a very important feature in rapid visual accommodation. Ciliary muscles change the shape of the lens to focus on near or far objects, a process known as accommodation. Predatory birds such as owls have long, tubular eyes and cannot readily accommodate their eyes to focus on very near prey. They often have to move away from their prey to better see it.
The vitreous body is a clear, jelly like substance that fills the eye between the lens and the retina. The viscosity of the vitreous supports and maintains the shape of the eye.
A small moveable membrane, the nictictating membrane (or third eyelid) is located in the corner of the eye. This membrane moves across the eye, spreading tear secretion produced by the gland of the nictitating membrane and a lacrinal gland. This tear secretion moistens the cornea and protects against micro organisms.
Winged Wisdom Note: Dr. Linda Pesek graduated from the University of Pennsylvania School of Veterinary Medicine. She has a small animal and avian practice in New York. Linda also writes columns for The Long Island Parrot Society and The Big Apple Bird Club and is a frequent lecturer at their meetings. She is the owner of an extensive collection of exotic birds.
Avian Reproduction: Anatomy & the Bird Egg THE BEST *****
Avian Reproductive System http://www.holisticbirds.com/pages/reproductive0803.htm
by Sherri Carpenter
The Avian Reproductive System
From cell to egg to a baby bird, is an amazing process. With the unity of a sperm cell to an ovum, a single cell develops into an embryo and a new life begins. I will begin with the reproductive organs.
Male Reproductive Organs
Male birds have a pair of testes that resemble a bean shape and each are located in front of the top lobe of the kidneys. During non-breeding season the testes are difficult to locate due to their small size, but during breeding season they may grow as much as several hundred times their non-breeding size.
As in mammals, the sperm cells of birds cannot develop fully at high temperatures that are found within the body cavity. Some birds experience a nightly drop in body temperature that allows the sperm cells to develop, while other birds have a swelling at the end of the tube (vas deferens). This tube connects the testes to the cloaca, and functions like a mammals scrotum holding the sperm away from the higher body temperatures that are within the abdomen.
Most bird species rub their cloacal areas together to transfer the male's sperm but ostriches, rheas, strokes, flamingos, ducks and a few other families actually have an erectile grooved penis on the back wall of the cloaca to transfer sperm.
Female Reproductive Organs
The female organs consist of the ovary and the oviduct that leads to the cloaca. With most bird species, the ovary is on the left side with the right side being underdeveloped and nonfunctional. It is thought that being only one sided reduces body weight and eliminates the possibility of carry two large, fragile eggs in the abdomen cavity at the same time.
The ovary when mature looks like a cluster of grapes. It may contain up to 4,000 small ova that can develop into yolks. Yolk protein, lipids and fats are manufactured in the liver and travel through the bloodstream to the immature ovum, during the maturation stage. Each yolk is attached to the ovary by a thin membrane sac or follicle having a fine network of blood vessels. The germinal disc of a developing yolk contains the single ovum cell which, after fertilization develops into a chick. The ovary enlarges during breeding season as much as fifty times its non-breeding weight.
The oviduct is a large coiled tube where all parts of the egg are formed except the yolk. It consists of several sections. Starting from the top: ostium, infundibular funnel, magnum area, isthmus, uterus and the vagina. They each play a part in the development of the egg.
When the brain’s pituitary gland releases a luteinizing hormone, LH, ovulation begins. The sac around the yolk ruptures and releases the yolk from the follicle, the yolk is kept intact by a fine membrane called the vitelline. The yolk is than engulfed by the infundibulum, with its thin, funnel-like lips. If the infundibulum cannot pick up the yolk, it is usually absorbed by the abdominal cavity or can cause peritonitis, an inflammtion of the lining of the abdominal cavity. Once inside the infundibulum fertilization can begin. The sperm have been stored in glands or nests, located in the infundibulum and are released as the yolk passes by. The sperm cell must penetrate the thin vitelline membrane and reach the female cell to complete fertilization.
The yolk than travel immediately into the magnum section for an stay of approximately three hour. In this time the egg white is added to the ovum. It is a protein substance containing mucin, globulin and albumen including sodium, magnesium and calcium. This serves as a shock absorber and feed the developing embryo.
In the isthmus the stay is approximately 75 minutes and another 10% of the albumen, the chalazae and shell membranes are added to the ovum. The chalazae are little ropes that attach to the yolk to keep it in the center and allow the yolk to rotate the keep the germinal disc ( the fertilized zygote) on top. The shell membranes are joined except in the area where the air cell will be, where they will separate.
The uterus is a thick-wall membrane where the egg will spend 20 hours. Through the shell membranes, water and salts are added which plump out the egg. The shell is added here and has three layers. The inner layer is first produced and called the mammillary layer. The middle layer is called testar and is the thickest and the outer layer is made of dried mucous. The shell is composed of calcium carbonate.
In the vagina the stay is short. A coating called “bloom” is added to keep harmful bacteria and or dust from entering the egg. The soft egg is shaped in the vagina depending on the shape of the bony pelvis.
The egg than travels to the cloaca and is laid. When the egg is laid, it is the same temperature as the hen and fills the shell. As it cools, it looses volume and the density changes slightly, creating a pressure, which draws air into the egg and forms the air cell. The shell hardens as it cools and dries. This whole process from start to finish has taken approximately 24 hours.
A brood spot develops on the hen and can develop on some males. The temperature rises on the brood spot due to a large amount of blood collecting in that area. Incubation of the eggs is usually 18 to 29 days depending on species. About 15% of an eggs original weight is lost due to evaporation in the incubation process. The eggs incubate at 99.1 to 99.5 degrees Fahrenheit with humidity between 50 and 53%. I will go into the embryo development in the next article.
Biology 325, Spring 1996
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