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126-724: Chronobiological studies include but are not limited to comparative anatomy , physiology , genetics , molecular biology and behavior of organisms related to their biological rhythms . Other aspects include epigenetics , development, reproduction, ecology and evolution. Chronobiology studies variations of the timing and duration of biological activity in living organisms which occur for many essential biological processes. These occur (a) in animals (eating, sleeping, mating, hibernating, migration, cellular regeneration, etc.), (b) in plants (leaf movements, photosynthetic reactions, etc.), and in microbial organisms such as fungi and protozoa. They have even been found in bacteria , especially among

252-412: A basal lamina , the upper layer of the basement membrane , the lower layer is the reticular lamina lying next to the connective tissue in the extracellular matrix secreted by the epithelial cells. There are many different types of epithelium, modified to suit a particular function. In the respiratory tract there is a type of ciliated epithelial lining; in the small intestine there are microvilli on

378-450: A copulatory organ present in most species. The eggs are surrounded by amniotic membranes which prevents them from drying out and are laid on land, or develop internally in some species. The bladder is small as nitrogenous waste is excreted as uric acid . Turtles are notable for their protective shells. They have an inflexible trunk encased in a horny carapace above and a plastron below. These are formed from bony plates embedded in

504-460: A flower clock using certain species of flowering plants . By arranging the selected species in a circular pattern, he designed a clock that indicated the time of day by the flowers that were open at each given hour. For example, among members of the daisy family , he used the hawk's beard plant which opened its flowers at 6:30 am and the hawkbit which did not open its flowers until 7 am. The 1960 symposium at Cold Spring Harbor Laboratory laid

630-481: A microscope . Human anatomy, physiology and biochemistry are complementary basic medical sciences, which are generally taught to medical students in their first year at medical school. Human anatomy can be taught regionally or systemically; that is, respectively, studying anatomy by bodily regions such as the head and chest, or studying by specific systems, such as the nervous or respiratory systems. The major anatomy textbook, Gray's Anatomy , has been reorganized from

756-470: A sessile lifestyle). Most animals have bodies differentiated into separate tissues and these animals are also known as eumetazoans . They have an internal digestive chamber, with one or two openings; the gametes are produced in multicellular sex organs, and the zygotes include a blastula stage in their embryonic development . Metazoans do not include the sponges , which have undifferentiated cells. Unlike plant cells , animal cells have neither

882-402: A "treatise on the heart", with vessels carrying all the body's fluids to or from every member of the body. Ancient Greek anatomy and physiology underwent great changes and advances throughout the early medieval world. Over time, this medical practice expanded due to a continually developing understanding of the functions of organs and structures in the body. Phenomenal anatomical observations of

1008-585: A T24 cycle, the T7 mice got the same amount of total sleep and their circadian expression of PER2 , an element of the SCN pacemaker, was not disrupted. Through the T7 cycle, the mice were exposed to light at all circadian phases. Light pulses presented at night lead to expression of the transcription factor c-Fos in the amygdala , lateral habenula , and subparaventricular nucleus further implicating light's possible influence on mood and other cognitive functions. Mice subjected to

1134-528: A cell wall nor chloroplasts . Vacuoles, when present, are more in number and much smaller than those in the plant cell. The body tissues are composed of numerous types of cells, including those found in muscles, nerves and skin. Each typically has a cell membrane formed of phospholipids , cytoplasm and a nucleus . All of the different cells of an animal are derived from the embryonic germ layers . Those simpler invertebrates which are formed from two germ layers of ectoderm and endoderm are called diploblastic and

1260-433: A class of arachnids have four pairs of legs; a body of two segments—a cephalothorax and an abdomen . Spiders have no wings and no antennae. They have mouthparts called chelicerae which are often connected to venom glands as most spiders are venomous. They have a second pair of appendages called pedipalps attached to the cephalothorax. These have similar segmentation to the legs and function as taste and smell organs. At

1386-552: A coat of glycoproteins . In more advanced animals, many glands are formed of epithelial cells. Muscle cells (myocytes) form the active contractile tissue of the body. Muscle tissue functions to produce force and cause motion, either locomotion or movement within internal organs. Muscle is formed of contractile filaments and is separated into three main types; smooth muscle , skeletal muscle and cardiac muscle . Smooth muscle has no striations when examined microscopically. It contracts slowly but maintains contractibility over

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1512-457: A few species have no limbs and resemble snakes. Lizards have moveable eyelids, eardrums are present and some species have a central parietal eye. Snakes are closely related to lizards, having branched off from a common ancestral lineage during the Cretaceous period, and they share many of the same features. The skeleton consists of a skull, a hyoid bone, spine and ribs though a few species retain

1638-449: A food-entrainable biological clock, which is not confined to the suprachiasmatic nucleus. The location of this clock has been disputed. Working with mice, however, Fuller et al. concluded that the food-entrainable clock seems to be located in the dorsomedial hypothalamus . During restricted feeding, it takes over control of such functions as activity timing, increasing the chances of the animal successfully locating food resources. In 2018

1764-513: A high metabolic rate , a light skeletal system and powerful muscles . The long bones are thin, hollow and very light. Air sac extensions from the lungs occupy the centre of some bones. The sternum is wide and usually has a keel and the caudal vertebrae are fused. There are no teeth and the narrow jaws are adapted into a horn-covered beak. The eyes are relatively large, particularly in nocturnal species such as owls. They face forwards in predators and sideways in ducks. The feathers are outgrowths of

1890-445: A key role in a variety of photic responses, including pupillary light reflex , and synchronization of the biological clock to daily light-dark cycles. He also described the role of melanopsin in ipRGCs . Using a rat melanopsin gene, a melanopsin-specific antibody, and fluorescent immunocytochemistry , the team concluded that melanopsin is expressed in some RGCs. Using a Beta-galactosidase assay, they found that these RGC axons exit

2016-406: A left Marcus Gunn pupil , or what is called left afferent pupillary defect, by physical examination. Location of the lesion can be deduced as follows: The pupillary response to light is not purely reflexive, but is modulated by cognitive factors, such as attention , awareness, and the way visual input is interpreted. For example, if a bright stimulus is presented to one eye, and a dark stimulus to

2142-746: A mammal. Humans have a head, neck, trunk (which includes the thorax and abdomen ), two arms and hands, and two legs and feet. Generally, students of certain biological sciences , paramedics , prosthetists and orthotists, physiotherapists , occupational therapists , nurses , podiatrists , and medical students learn gross anatomy and microscopic anatomy from anatomical models, skeletons, textbooks, diagrams, photographs, lectures and tutorials and in addition, medical students generally also learn gross anatomy through practical experience of dissection and inspection of cadavers . The study of microscopic anatomy (or histology ) can be aided by practical experience examining histological preparations (or slides) under

2268-498: A means to reset animal and human circadian rhythms. Additionally, the presence of low-level light at night accelerates circadian re-entrainment of hamsters of all ages by 50%; this is thought to be related to simulation of moonlight. In the second half of 20th century, substantial contributions and formalizations have been made by Europeans such as Jürgen Aschoff and Colin Pittendrigh , who pursued different but complementary views on

2394-464: A number of central targets. Five classes of ipRGCs, M1 through M5, have been characterized to date in rodents. These classes differ in morphology, dendritic localization, melanopsin content, electrophysiological profiles, and projections. Hattar and his co-workers discovered that, even among the subtypes of ipRGC, there can be designated sets that differentially control circadian versus pupillary behavior. In experiments with M1 ipRGCs, they discovered that

2520-420: A pump action in which air is first drawn into the buccopharyngeal region through the nostrils. These are then closed and the air is forced into the lungs by contraction of the throat. They supplement this with gas exchange through the skin which needs to be kept moist. In frogs the pelvic girdle is robust and the hind legs are much longer and stronger than the forelimbs. The feet have four or five digits and

2646-421: A role in dilating the pupils in low light conditions. The optic nerve , or more precisely, the photosensitive ganglion cells through the retinohypothalamic tract , is responsible for the afferent limb of the pupillary reflex; it senses the incoming light. The oculomotor nerve is responsible for the efferent limb of the pupillary reflex; it drives the iris muscles that constrict the pupil. Referring to

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2772-400: A significant increase in the use of advanced imaging techniques, such as MRI and CT scans , which allow for more detailed and accurate visualizations of the body's structures. The discipline of anatomy is divided into macroscopic and microscopic parts. Macroscopic anatomy , or gross anatomy, is the examination of an animal's body parts using unaided eyesight. Gross anatomy also includes

2898-543: A skeleton to support or protect the body. An exoskeleton is a thickened, rigid cuticle which is stiffened by mineralization , as in crustaceans or by the cross-linking of its proteins as in insects . An endoskeleton is internal and present in all developed animals, as well as in many of those less developed. Epithelial tissue is composed of closely packed cells, bound to each other by cell adhesion molecules , with little intercellular space. Epithelial cells can be squamous (flat), cuboidal or columnar and rest on

3024-572: A study published in PLoS ONE showed how 73 psychometric indicators measured on Twitter Content follow a diurnal pattern. A followup study appeared on Chronobiology International in 2021 showed that these patterns were not disrupted by the 2020 UK lockdown. In 2021, scientists reported the development of a light-responsive days-lasting modulator of circadian rhythms of tissues via Ck1 inhibition . Such modulators may be useful for chronobiology research and repair of organs that are "out of sync". Chronobiology

3150-497: A systems format to a regional format, in line with modern teaching methods. A thorough working knowledge of anatomy is required by physicians, especially surgeons and doctors working in some diagnostic specialties, such as histopathology and radiology . Academic anatomists are usually employed by universities, medical schools or teaching hospitals. They are often involved in teaching anatomy, and research into certain systems, organs, tissues or cells. Invertebrates constitute

3276-431: A vast array of living organisms ranging from the simplest unicellular eukaryotes such as Paramecium to such complex multicellular animals as the octopus , lobster and dragonfly . They constitute about 95% of the animal species. By definition, none of these creatures has a backbone. The cells of single-cell protozoans have the same basic structure as those of multicellular animals but some parts are specialized into

3402-479: A vertebrate is the vertebral column , formed in the development of the segmented series of vertebrae . In most vertebrates the notochord becomes the nucleus pulposus of the intervertebral discs . However, a few vertebrates, such as the sturgeon and the coelacanth , retain the notochord into adulthood. Jawed vertebrates are typified by paired appendages, fins or legs, which may be secondarily lost. The limbs of vertebrates are considered to be homologous because

3528-693: A vestige of the pelvis and rear limbs in the form of pelvic spurs . The bar under the second fenestra has also been lost and the jaws have extreme flexibility allowing the snake to swallow its prey whole. Snakes lack moveable eyelids, the eyes being covered by transparent "spectacle" scales. They do not have eardrums but can detect ground vibrations through the bones of their skull. Their forked tongues are used as organs of taste and smell and some species have sensory pits on their heads enabling them to locate warm-blooded prey. Crocodilians are large, low-slung aquatic reptiles with long snouts and large numbers of teeth. The head and trunk are dorso-ventrally flattened and

3654-594: A wide range of stretch lengths. It is found in such organs as sea anemone tentacles and the body wall of sea cucumbers . Skeletal muscle contracts rapidly but has a limited range of extension. It is found in the movement of appendages and jaws. Obliquely striated muscle is intermediate between the other two. The filaments are staggered and this is the type of muscle found in earthworms that can extend slowly or make rapid contractions. In higher animals striated muscles occur in bundles attached to bone to provide movement and are often arranged in antagonistic sets. Smooth muscle

3780-437: Is a branch of natural science that deals with the structural organization of living things. It is an old science, having its beginnings in prehistoric times. Anatomy is inherently tied to developmental biology , embryology , comparative anatomy , evolutionary biology , and phylogeny , as these are the processes by which anatomy is generated, both over immediate and long-term timescales. Anatomy and physiology , which study

3906-451: Is an interdisciplinary field of investigation. It interacts with medical and other research fields such as sleep medicine , endocrinology , geriatrics , sports medicine , space medicine and photoperiodism . Anatomy Anatomy (from Ancient Greek ἀνατομή ( anatomḗ )  ' dissection ') is the branch of morphology concerned with the study of the internal structure of organisms and their parts. Anatomy

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4032-473: Is found in the walls of the uterus , bladder , intestines , stomach , oesophagus , respiratory airways , and blood vessels . Cardiac muscle is found only in the heart , allowing it to contract and pump blood round the body. Nervous tissue is composed of many nerve cells known as neurons which transmit information. In some slow-moving radially symmetrical marine animals such as ctenophores and cnidarians (including sea anemones and jellyfish ),

4158-463: Is modulated by subjective (as opposed to objective) brightness. Pupillary light reflex is modeled as a physiologically-based non-linear delay differential equation that describes the changes in the pupil diameter as a function of the environment lighting: where D {\displaystyle D} is the pupil diameter measured in millimeters and Φ ( t − τ ) {\displaystyle \Phi (t-\tau )}

4284-402: Is modulated by visual attention and trial-by-trial variation in visual attention. Finally, a picture that is subjectively perceived as bright (e.g. a picture of the sun), elicits a stronger pupillary constriction than an image that is perceived as less bright (e.g. a picture of an indoor scene), even when the objective brightness of both images is equal. This shows that the pupillary light reflex

4410-464: Is opened and its organs studied, and endoscopy , in which a video camera -equipped instrument is inserted through a small incision in the body wall and used to explore the internal organs and other structures. Angiography using X-rays or magnetic resonance angiography are methods to visualize blood vessels. The term "anatomy" is commonly taken to refer to human anatomy . However, substantially similar structures and tissues are found throughout

4536-489: Is startlingly rapid". Between 1275 and 1326, the anatomists Mondino de Luzzi , Alessandro Achillini and Antonio Benivieni at Bologna carried out the first systematic human dissections since ancient times. Mondino's Anatomy of 1316 was the first textbook in the medieval rediscovery of human anatomy. It describes the body in the order followed in Mondino's dissections, starting with the abdomen, thorax, head, and limbs. It

4662-407: Is synonymous with pupillary response, which may be pupillary constriction or dilation. Pupillary reflex is conceptually linked to the side (left or right) of the reacting pupil, and not to the side from which light stimulation originates. Left pupillary reflex refers to the response of the left pupil to light, regardless of which eye is exposed to a light source. Right pupillary reflex means reaction of

4788-401: Is the luminous intensity reaching the retina in a time t {\displaystyle t} , which can be described as Φ = I A {\displaystyle \Phi =IA} : luminance reaching the eye in lumens/mm times the pupil area in mm . τ {\displaystyle \tau } is the pupillary latency, a time delay between the instant in which

4914-403: Is the study of the structures that make up a discrete body system—that is, a group of structures that work together to perform a unique body function, such as the digestive system. Anatomy can be studied using both invasive and non-invasive methods with the goal of obtaining information about the structure and organization of organs and systems. Methods used include dissection , in which a body

5040-411: Is the vertebral column, composed of articulating vertebrae which are lightweight yet strong. The ribs attach to the spine and there are no limbs or limb girdles. The main external features of the fish, the fins , are composed of either bony or soft spines called rays, which with the exception of the caudal fins , have no direct connection with the spine. They are supported by the muscles which compose

5166-590: The Hippocratic Corpus , an Ancient Greek medical work written by unknown authors. Aristotle described vertebrate anatomy based on animal dissection . Praxagoras identified the difference between arteries and veins . Also in the 4th century BCE, Herophilos and Erasistratus produced more accurate anatomical descriptions based on vivisection of criminals in Alexandria during the Ptolemaic period . In

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5292-677: The Society for Research in Biological Rhythms in the 1970s. Halberg wanted more emphasis on the human and medical issues while Pittendrigh had his background more in evolution and ecology. With Pittendrigh as leader, the Society members did basic research on all types of organisms, plants as well as animals. More recently it has been difficult to get funding for such research on any other organisms than mice, rats, humans and fruit flies . In 2002, Hattar and his colleagues showed that melanopsin plays

5418-506: The Triassic period. There is one living species, Sphenodon punctatus . The skull has two openings (fenestrae) on either side and the jaw is rigidly attached to the skull. There is one row of teeth in the lower jaw and this fits between the two rows in the upper jaw when the animal chews. The teeth are merely projections of bony material from the jaw and eventually wear down. The brain and heart are more primitive than those of other reptiles, and

5544-510: The cyanobacteria (aka blue-green algae, see bacterial circadian rhythms ). The best studied rhythm in chronobiology is the circadian rhythm , a roughly 24-hour cycle shown by physiological processes in all these organisms. The term circadian comes from the Latin circa , meaning "around" and dies , "day", meaning "approximately a day." It is regulated by circadian clocks . The circadian rhythm can further be broken down into routine cycles during

5670-400: The epidermis and are found in localized bands from where they fan out over the skin. Large flight feathers are found on the wings and tail, contour feathers cover the bird's surface and fine down occurs on young birds and under the contour feathers of water birds. The only cutaneous gland is the single uropygial gland near the base of the tail. This produces an oily secretion that waterproofs

5796-529: The extracellular matrix . Often called fascia (from the Latin "fascia," meaning "band" or "bandage"), connective tissues give shape to organs and holds them in place. The main types are loose connective tissue, adipose tissue , fibrous connective tissue, cartilage and bone. The extracellular matrix contains proteins , the chief and most abundant of which is collagen . Collagen plays a major part in organizing and maintaining tissues. The matrix can be modified to form

5922-472: The iris and is where light enters the eye. By analogy with a camera, the pupil is equivalent to aperture , whereas the iris is equivalent to the diaphragm . It may be helpful to consider the Pupillary reflex as an ' Iris' reflex , as the iris sphincter and dilator muscles are what can be seen responding to ambient light. Whereas, the pupil is the passive opening formed by the active iris. Pupillary reflex

6048-461: The mouthparts . The thorax has three pairs of segmented legs , one pair each for the three segments that compose the thorax and one or two pairs of wings . The abdomen is composed of eleven segments, some of which may be fused and houses the digestive , respiratory , excretory and reproductive systems. There is considerable variation between species and many adaptations to the body parts, especially wings, legs, antennae and mouthparts. Spiders

6174-400: The stomach . All vertebrates have a similar basic body plan and at some point in their lives, mostly in the embryonic stage, share the major chordate characteristics: a stiffening rod, the notochord ; a dorsal hollow tube of nervous material, the neural tube ; pharyngeal arches ; and a tail posterior to the anus. The spinal cord is protected by the vertebral column and is above

6300-543: The 24-hour day: While circadian rhythms are defined as regulated by endogenous processes, other biological cycles may be regulated by exogenous signals. In some cases, multi-trophic systems may exhibit rhythms driven by the circadian clock of one of the members (which may also be influenced or reset by external factors). The endogenous plant cycles may regulate the activity of the bacterium by controlling availability of plant-produced photosynthate. Many other important cycles are also studied, including: Within each cycle,

6426-420: The 2nd century, Galen of Pergamum , an anatomist, clinician , writer, and philosopher, wrote the final and highly influential anatomy treatise of ancient times. He compiled existing knowledge and studied anatomy through the dissection of animals. He was one of the first experimental physiologists through his vivisection experiments on animals. Galen's drawings, based mostly on dog anatomy, became effectively

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6552-462: The Edinger-Westphal nucleus on the contralateral side. Segments 5 and 6 are fibers that connect the pretectal nucleus on one side to the Edinger-Westphal nucleus on the same side. Segments 3, 4, 5, and 6 are all located within a compact region within the midbrain. Segments 7 and 8 each contains parasympathetic fibers that courses from the Edinger-Westphal nucleus, through the ciliary ganglion, along

6678-497: The M1 ipRGC consist of molecularly distinct subpopulations that innervate different brain regions and execute specific light-induced functions. This isolation of a 'labeled line' consisting of differing molecular and functional properties in a highly specific ipRGC subtype was an important first for the field. It also underscored the extent to which molecular signatures can be used to distinguish between RGC populations that would otherwise appear

6804-758: The SCN, where input from ipRGCs is necessary to photoentrain circadian rhythms, and the olivary pretectal nucleus (OPN), where input from ipRGCs control the pupillary light reflex. Hattar and colleagues conducted research that demonstrated that ipRGCs project to hypothalamic, thalamic, stratal, brainstem and limbic structures. Although ipRGCs were initially viewed as a uniform population, further research revealed that there are several subtypes with distinct morphology and physiology. Since 2011, Hattar's laboratory has contributed to these findings and has successfully distinguished subtypes of ipRGCs. Hattar and colleges utilized Cre-based strategies for labeling ipRGCs to reveal that there are at least five ipRGC subtypes that project to

6930-454: The T7 cycle exhibited depression-like symptoms, exhibiting decreased preference for sucrose (sucrose anhedonia) and exhibiting more immobility than their T24 counterparts in the forced swim test (FST) . Additionally, T7 mice maintained rhythmicity in serum corticosterone , however the levels were elevated compared to the T24 mice, a trend that is associated with depression. Chronic administration of

7056-604: The T7 mice in the Morris water maze (MWM) , a spatial learning task that places a mouse in a small pool of water and tests the mouse's ability to locate and remember the location of a rescue platform located just below the waterline. Compared to the T24 mice, the T7 mice took longer to find the platform in subsequent trials and did not exhibit a preference for the quadrant containing the platform. In addition, T7 mice exhibited impaired hippocampal long-term potentiation (LTP) when subjected to theta burst stimulation (TBS). Recognition memory

7182-517: The antidepressant Fluoxetine lowered corticosterone levels in T7 mice and reduced depression-like behavior while leaving their circadian rhythms unaffected. The hippocampus is a structure in the limbic system that receives projections from ipRGCs. It is required for the consolidation of short-term memories into long-term memories as well as spatial orientation and navigation. Depression and heightened serum corticosterone levels are linked to impaired hippocampal learning. Hattar and his team analyzed

7308-425: The appearance and position of the various parts, the materials from which they are composed, and their relationships with other parts. Anatomy is quite distinct from physiology and biochemistry , which deal respectively with the functions of those parts and the chemical processes involved. For example, an anatomist is concerned with the shape, size, position, structure, blood supply and innervation of an organ such as

7434-531: The arts and sciences from the Ptolemaic dynasty of Egypt helped raise Alexandria up, further rivalling other Greek states' cultural and scientific achievements. Some of the most striking advances in early anatomy and physiology took place in Hellenistic Alexandria. Two of the most famous anatomists and physiologists of the third century were Herophilus and Erasistratus . These two physicians helped pioneer human dissection for medical research, using

7560-490: The atria were parts of the heart. Herophilus's knowledge of the human body has provided vital input towards understanding the brain, eye, liver, reproductive organs, and nervous system and characterizing the course of the disease. Erasistratus accurately described the structure of the brain, including the cavities and membranes, and made a distinction between its cerebrum and cerebellum During his study in Alexandria, Erasistratus

7686-518: The basis of sense organs and there is a central nervous system (brain and spinal cord) and a peripheral nervous system . The latter consists of sensory nerves that transmit information from sense organs and motor nerves that influence target organs. The peripheral nervous system is divided into the somatic nervous system which conveys sensation and controls voluntary muscle , and the autonomic nervous system which involuntarily controls smooth muscle , certain glands and internal organs, including

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7812-462: The body and they swim by undulating their body from side to side. Reptiles are a class of animals comprising turtles , tuataras , lizards , snakes and crocodiles . They are tetrapods , but the snakes and a few species of lizard either have no limbs or their limbs are much reduced in size. Their bones are better ossified and their skeletons stronger than those of amphibians. The teeth are conical and mostly uniform in size. The surface cells of

7938-619: The brain that are important for regulating circadian rhythmicity and sleep, most notably the SCN , subparaventricular nucleus, and the ventrolateral preoptic area. In addition, ipRGCs transmit information to many areas in the limbic system , which is strongly tied to emotion and memory. To examine the relationship between deviant light exposure and behavior, Hattar and his colleagues studied mice exposed to alternating 3.5-hour light and dark periods (T7 mice) and compared them with mice exposed to alternating 12-hour light and dark periods (T24 mice). Compared to

8064-399: The brain, appreciated the division between cerebellum and cerebrum and recognized that the brain was the "seat of intellect" and not a "cooling chamber" as propounded by Aristotle Herophilus is also credited with describing the optic, oculomotor, motor division of the trigeminal, facial, vestibulocochlear and hypoglossal nerves. Incredible feats were made during the third century BCE in both

8190-401: The branch of superficial anatomy . Microscopic anatomy involves the use of optical instruments in the study of the tissues of various structures, known as histology , and also in the study of cells . The history of anatomy is characterized by a progressive understanding of the functions of the organs and structures of the human body. Methods have also improved dramatically, advancing from

8316-516: The cadavers of condemned criminals, which was considered taboo until the Renaissance—Herophilus was recognized as the first person to perform systematic dissections. Herophilus became known for his anatomical works, making impressive contributions to many branches of anatomy and many other aspects of medicine. Some of the works included classifying the system of the pulse, the discovery that human arteries had thicker walls than veins, and that

8442-427: The ciliary ganglion. Postganglionic nerve fibers leave the ciliary ganglion to innervate the pupillary sphincter. Each afferent limb has two efferent limbs, one ipsilateral and one contralateral. The ipsilateral efferent limb transmits nerve signals for direct light reflex of the ipsilateral pupil. The contralateral efferent limb causes consensual light reflex of the contralateral pupil. Sympathetic nervous system plays

8568-454: The current and previous simulation times (times since the simulation started) measured in milliseconds, S {\displaystyle S} is a constant that affects the constriction/dilation velocity and varies among individuals. The higher the S {\displaystyle S} value, the smaller the time step used in the simulation and, consequently, the smaller the pupil constriction/dilation velocity. In order to improve

8694-418: The dermis which are overlain by horny ones and are partially fused with the ribs and spine. The neck is long and flexible and the head and the legs can be drawn back inside the shell. Turtles are vegetarians and the typical reptile teeth have been replaced by sharp, horny plates. In aquatic species, the front legs are modified into flippers. Tuataras superficially resemble lizards but the lineages diverged in

8820-402: The digestive and reproductive systems. Herophilus discovered and described not only the salivary glands but also the small intestine and liver. He showed that the uterus is a hollow organ and described the ovaries and uterine tubes. He recognized that spermatozoa were produced by the testes and was the first to identify the prostate gland. The anatomy of the muscles and skeleton is described in

8946-402: The egg-laying monotremes , the platypus and the echidnas of Australia. Most other mammals have a placenta through which the developing foetus obtains nourishment, but in marsupials , the foetal stage is very short and the immature young is born and finds its way to its mother's pouch where it latches on to a teat and completes its development. Humans have the overall body plan of

9072-523: The end of each male pedipalp is a spoon-shaped cymbium that acts to support the copulatory organ . In 1600 BCE, the Edwin Smith Papyrus , an Ancient Egyptian medical text , described the heart and its vessels, as well as the brain and its meninges and cerebrospinal fluid , and the liver , spleen , kidneys , uterus and bladder . It showed the blood vessels diverging from the heart. The Ebers Papyrus ( c.  1550 BCE ) features

9198-467: The epidermis are modified into horny scales which create a waterproof layer. Reptiles are unable to use their skin for respiration as do amphibians and have a more efficient respiratory system drawing air into their lungs by expanding their chest walls. The heart resembles that of the amphibian but there is a septum which more completely separates the oxygenated and deoxygenated bloodstreams. The reproductive system has evolved for internal fertilization, with

9324-413: The epidermis is normally formed of epithelial cells and secretes an extracellular matrix which provides support to the organism. An endoskeleton derived from the mesoderm is present in echinoderms , sponges and some cephalopods . Exoskeletons are derived from the epidermis and is composed of chitin in arthropods (insects, spiders, ticks, shrimps, crabs, lobsters). Calcium carbonate constitutes

9450-429: The epithelial lining and in the large intestine there are intestinal villi . Skin consists of an outer layer of keratinized stratified squamous epithelium that covers the exterior of the vertebrate body. Keratinocytes make up to 95% of the cells in the skin . The epithelial cells on the external surface of the body typically secrete an extracellular matrix in the form of a cuticle . In simple animals this may just be

9576-610: The equivalent of tissues and organs. Locomotion is often provided by cilia or flagella or may proceed via the advance of pseudopodia , food may be gathered by phagocytosis , energy needs may be supplied by photosynthesis and the cell may be supported by an endoskeleton or an exoskeleton . Some protozoans can form multicellular colonies. Metazoans are a multicellular organism, with different groups of cells serving different functions. The most basic types of metazoan tissues are epithelium and connective tissue, both of which are present in nearly all invertebrates. The outer surface of

9702-527: The examination of animals by dissection of carcasses and cadavers (corpses) to 20th-century medical imaging techniques, including X-ray , ultrasound , and magnetic resonance imaging . Derived from the Greek ἀνατομή anatomē "dissection" (from ἀνατέμνω anatémnō "I cut up, cut open" from ἀνά aná "up", and τέμνω témnō "I cut"), anatomy is the scientific study of the structure of organisms including their systems, organs and tissues . It includes

9828-499: The eye. Emergency physicians routinely test pupillary light reflex to assess brain stem function. Abnormal pupillary reflex can be found in optic nerve injury, oculomotor nerve damage, brain stem lesion (including brain stem death ), and depressant drugs, such as barbiturates . Examples are provided as below: For example, in a person with abnormal left direct reflex and abnormal right consensual reflex (with normal left consensual and normal right direct reflexes), which would produce

9954-554: The eyes together with the optic nerve and project to the suprachiasmatic nucleus (SCN), the primary circadian pacemaker in mammals . They also demonstrated that the RGCs containing melanopsin were intrinsically photosensitive. Hattar concluded that melanopsin is the photopigment in a small subset of RGCs that contributes to the intrinsic photosensitivity of these cells and is involved in their non-image forming functions, such as photic entrainment and pupillary light reflex. Hattar, armed with

10080-405: The father of microscopical anatomy, discovered that plants had tubules similar to those he saw in insects like the silk worm. He observed that when a ring-like portion of bark was removed on a trunk a swelling occurred in the tissues above the ring, and he unmistakably interpreted this as growth stimulated by food coming down from the leaves, and being captured above the ring. Arthropods comprise

10206-435: The feathers when the bird preens . There are scales on the legs, feet and claws on the tips of the toes. Mammals are a diverse class of animals, mostly terrestrial but some are aquatic and others have evolved flapping or gliding flight. They mostly have four limbs, but some aquatic mammals have no limbs or limbs modified into fins, and the forelimbs of bats are modified into wings. The legs of most mammals are situated below

10332-443: The features of ancient fish. They have a bony skeleton, are generally laterally flattened, have five pairs of gills protected by an operculum , and a mouth at or near the tip of the snout. The dermis is covered with overlapping scales . Bony fish have a swim bladder which helps them maintain a constant depth in the water column, but not a cloaca. They mostly spawn a large number of small eggs with little yolk which they broadcast into

10458-453: The groundwork for the field of chronobiology. It was also in 1960 that Patricia DeCoursey invented the phase response curve , one of the major tools used in the field since. Franz Halberg of the University of Minnesota , who coined the word circadian , is widely considered the "father of American chronobiology." However, it was Colin Pittendrigh and not Halberg who was elected to lead

10584-482: The head. The dermis is covered with separate dermal placoid scales . They have a cloaca into which the urinary and genital passages open, but not a swim bladder . Cartilaginous fish produce a small number of large, yolky eggs. Some species are ovoviviparous and the young develop internally but others are oviparous and the larvae develop externally in egg cases. The bony fish lineage shows more derived anatomical traits, often with major evolutionary changes from

10710-512: The human body were made, which contributed to the understanding of the brain, eye, liver, reproductive organs, and nervous system. The Hellenistic Egyptian city of Alexandria was the stepping-stone for Greek anatomy and physiology. Alexandria not only housed the biggest library for medical records and books of the liberal arts in the world during the time of the Greeks but was also home to many medical practitioners and philosophers. Great patronage of

10836-529: The ipsilateral (same) eye. A consensual pupillary reflex is response of a pupil to light that enters the contralateral (opposite) eye. Thus there are four types of pupillary light reflexes, based on this terminology of absolute laterality (left versus right) and relative laterality (same side versus opposite side, ipsilateral versus contralateral, direct versus consensual): The pupillary light reflex neural pathway on each side has an afferent limb and two efferent limbs. The afferent limb has nerve fibers running within

10962-460: The ipsilateral pretectal nucleus. The efferent limb is the pupillary motor output from the pretectal nucleus to the pupillary sphincter of the iris. The pretectal nucleus projects nerve fibers to the ipsilateral and contralateral Edinger-Westphal nuclei , which are also located in the midbrain. Each Edinger-Westphal nucleus gives rise to preganglionic parasympathetic fibers which exit with CN III and synapse with postganglionic parasympathetic neurons in

11088-813: The knowledge that melanopsin was the photopigment responsible for the photosensitivity of ipRGCs, set out to study the exact role of the ipRGC in photoentrainment . In 2008, Hattar and his research team transplanted diphtheria toxin genes into the mouse melanopsin gene locus to create mutant mice that lacked ipRGCs. The research team found that while the mutants had little difficulty identifying visual targets, they could not entrain to light-dark cycles. These results led Hattar and his team to conclude that ipRGCs do not affect image-forming vision, but significantly affect non-image forming functions such as photoentrainment. Further research has shown that ipRGCs project to different brain nuclei to control both non-image forming and image forming functions. These brain regions include

11214-492: The largest phylum of invertebrates in the animal kingdom with over a million known species. Insects possess segmented bodies supported by a hard-jointed outer covering, the exoskeleton , made mostly of chitin . The segments of the body are organized into three distinct parts, a head, a thorax and an abdomen . The head typically bears a pair of sensory antennae , a pair of compound eyes , one to three simple eyes ( ocelli ) and three sets of modified appendages that form

11340-399: The length of the sides of fish, and these respond to nearby movements and to changes in water pressure. Sharks and rays are basal fish with numerous primitive anatomical features similar to those of ancient fish, including skeletons composed of cartilage. Their bodies tend to be dorso-ventrally flattened, they usually have five pairs of gill slits and a large mouth set on the underside of

11466-559: The light pulse reaches the retina and the beginning of iridal reaction due nerve transmission, neuro-muscular excitation and activation delays. d M {\displaystyle \mathrm {d} M} , d D {\displaystyle \mathrm {d} D} and d t {\displaystyle \mathrm {d} t} are the derivatives for the M {\displaystyle M} function, pupil diameter D {\displaystyle D} and time t {\displaystyle t} . Since

11592-439: The liver; while a physiologist is interested in the production of bile , the role of the liver in nutrition and the regulation of bodily functions. The discipline of anatomy can be subdivided into a number of branches, including gross or macroscopic anatomy and microscopic anatomy. Gross anatomy is the study of structures large enough to be seen with the naked eye, and also includes superficial anatomy or surface anatomy,

11718-437: The lungs have a single chamber and lack bronchi . The tuatara has a well-developed parietal eye on its forehead. Lizards have skulls with only one fenestra on each side, the lower bar of bone below the second fenestra having been lost. This results in the jaws being less rigidly attached which allows the mouth to open wider. Lizards are mostly quadrupeds, with the trunk held off the ground by short, sideways-facing legs, but

11844-400: The main part of the trunk. The heart has two chambers and pumps the blood through the respiratory surfaces of the gills and on round the body in a single circulatory loop. The eyes are adapted for seeing underwater and have only local vision. There is an inner ear but no external or middle ear . Low frequency vibrations are detected by the lateral line system of sense organs that run along

11970-477: The more developed animals whose structures and organs are formed from three germ layers are called triploblastic . All of a triploblastic animal's tissues and organs are derived from the three germ layers of the embryo, the ectoderm , mesoderm and endoderm . Animal tissues can be grouped into four basic types: connective , epithelial , muscle and nervous tissue . Connective tissues are fibrous and made up of cells scattered among inorganic material called

12096-475: The nerves form a nerve net , but in most animals they are organized longitudinally into bundles. In simple animals, receptor neurons in the body wall cause a local reaction to a stimulus. In more complex animals, specialized receptor cells such as chemoreceptors and photoreceptors are found in groups and send messages along neural networks to other parts of the organism. Neurons can be connected together in ganglia . In higher animals, specialized receptors are

12222-435: The neural pathway schematic diagram, the entire pupillary light reflex system can be visualized as having eight neural segments, numbered 1 through 8. Odd-numbered segments 1, 3, 5, and 7 are on the left. Even-numbered segments 2, 4, 6, and 8 are on the right. Segments 1 and 2 each includes both the retina and the optic nerve (cranial Nerve #2). Segments 3 and 4 are nerve fibers that cross from the pretectal nucleus on one side to

12348-416: The notochord, and the gastrointestinal tract is below it. Nervous tissue is derived from the ectoderm , connective tissues are derived from mesoderm , and gut is derived from the endoderm . At the posterior end is a tail which continues the spinal cord and vertebrae but not the gut. The mouth is found at the anterior end of the animal, and the anus at the base of the tail. The defining characteristic of

12474-410: The oculomotor nerve (cranial nerve #3), to the ciliary sphincter, the muscular structure within the iris. The diagram may assist in localizing lesion within the pupillary reflex system by process of elimination, using light reflex testing results obtained by clinical examination. Pupillary light reflex provides a useful diagnostic tool for testing the integrity of the sensory and motor functions of

12600-428: The only anatomical textbook for the next thousand years. His work was known to Renaissance doctors only through Islamic Golden Age medicine until it was translated from Greek sometime in the 15th century. Anatomy developed little from classical times until the sixteenth century; as the historian Marie Boas writes, "Progress in anatomy before the sixteenth century is as mysteriously slow as its development after 1500

12726-407: The optic nerve ( CN II ). Each efferent limb has parasympathetic nerve fibers traveling along the periphery of the oculomotor nerve ( CN III ). The afferent limb carries sensory input. Anatomically, the afferent limb consists of the retina, the optic nerve, and the pretectal nucleus in the midbrain, at level of superior colliculus. Ganglion cells of the retina project fibers through the optic nerve to

12852-464: The other eye, perception alternates between the two eyes (i.e., binocular rivalry ): Sometimes the dark stimulus is perceived, sometimes the bright stimulus, but never both at the same time. Using this technique, it has been shown the pupil is smaller when a bright stimulus dominates awareness, relative to when a dark stimulus dominates awareness. This shows that the pupillary light reflex is modulated by visual awareness. Similarly, it has been shown that

12978-402: The phenomenon of entrainment of the circadian system by light (parametric, continuous, tonic, gradual vs. nonparametric, discrete, phasic, instantaneous, respectively). Humans can have a propensity to be morning people or evening people; these behavioral preferences are called chronotypes for which there are various assessment questionnaires and biological marker correlations. There is also

13104-633: The pupil constriction velocity is approximately 3 times faster than (re)dilation velocity, different step sizes in the numerical solver simulation must be used: where d t c {\displaystyle \mathrm {d} t_{c}} and d t d {\displaystyle \mathrm {d} t_{d}} are respectively the d t {\displaystyle \mathrm {d} t} for constriction and dilation measured in milliseconds, T c {\displaystyle T_{c}} and T p {\displaystyle T_{p}} are respectively

13230-420: The pupil constricts when you covertly (i.e., without looking at) pay attention to a bright stimulus, compared to a dark stimulus, even when visual input is identical. Moreover, the magnitude of the pupillary light reflex following a distracting probe is strongly correlated with the extent to which the probe captures visual attention and interferes with task performance. This shows that the pupillary light reflex

13356-399: The pupil to constrict ( miosis/myosis ; thereby allowing less light in), whereas a lower intensity of light causes the pupil to dilate ( mydriasis , expansion; thereby allowing more light in). Thus, the pupillary light reflex regulates the intensity of light entering the eye. Light shone into one eye will cause both pupils to constrict. The pupil is the dark circular opening in the center of

13482-419: The rest of the animal kingdom, and the term also includes the anatomy of other animals. The term zootomy is also sometimes used to specifically refer to non-human animals. The structure and tissues of plants are of a dissimilar nature and they are studied in plant anatomy . The kingdom Animalia contains multicellular organisms that are heterotrophic and motile (although some have secondarily adopted

13608-401: The right pupil, whether light is shone into the left eye, right eye, or both eyes. When light is shone into only one eye and not the other, it is normal for both pupils to constrict simultaneously. The terms direct and consensual refers to the side where the light source comes from, relative to the side of the reacting pupil. A direct pupillary reflex is pupillary response to light that enters

13734-493: The same underlying skeletal structure was inherited from their last common ancestor. This is one of the arguments put forward by Charles Darwin to support his theory of evolution . The body of a fish is divided into a head, trunk and tail, although the divisions between the three are not always externally visible. The skeleton, which forms the support structure inside the fish, is either made of cartilage, in cartilaginous fish , or bone in bony fish . The main skeletal element

13860-440: The same, which in turn facilitates further investigation into their specific contributions to visual processing. Previous studies in circadian biology have established that exposure to light during abnormal hours leads to sleep deprivation and disruption of the circadian system, which affect mood and cognitive functioning . While this indirect relationship had been corroborated, not much work had been done to examine whether there

13986-670: The shells of molluscs , brachiopods and some tube-building polychaete worms and silica forms the exoskeleton of the microscopic diatoms and radiolaria . Other invertebrates may have no rigid structures but the epidermis may secrete a variety of surface coatings such as the pinacoderm of sponges, the gelatinous cuticle of cnidarians ( polyps , sea anemones , jellyfish ) and the collagenous cuticle of annelids . The outer epithelial layer may include cells of several types including sensory cells, gland cells and stinging cells. There may also be protrusions such as microvilli , cilia, bristles, spines and tubercles . Marcello Malpighi ,

14112-404: The structure and function of organisms and their parts respectively, make a natural pair of related disciplines, and are often studied together. Human anatomy is one of the essential basic sciences that are applied in medicine, and is often studied alongside physiology . Anatomy is a complex and dynamic field that is constantly evolving as discoveries are made. In recent years, there has been

14238-400: The study by sight of the external body features. Microscopic anatomy is the study of structures on a microscopic scale, along with histology (the study of tissues), and embryology (the study of an organism in its immature condition). Regional anatomy is the study of the interrelationships of all of the structures in a specific body region, such as the abdomen. In contrast, systemic anatomy

14364-741: The tail is laterally compressed. It undulates from side to side to force the animal through the water when swimming. The tough keratinized scales provide body armour and some are fused to the skull. The nostrils, eyes and ears are elevated above the top of the flat head enabling them to remain above the surface of the water when the animal is floating. Valves seal the nostrils and ears when it is submerged. Unlike other reptiles, crocodilians have hearts with four chambers allowing complete separation of oxygenated and deoxygenated blood. Birds are tetrapods but though their hind limbs are used for walking or hopping, their front limbs are wings covered with feathers and adapted for flight. Birds are endothermic , have

14490-561: The time period during which the process is more active is called the acrophase . When the process is less active, the cycle is in its bathyphase or trough phase. The particular moment of highest activity is the peak or maximum ; the lowest point is the nadir . A circadian cycle was first observed in the 18th century in the movement of plant leaves by the French scientist Jean-Jacques d'Ortous de Mairan . In 1751 Swedish botanist and naturalist Carl Linnaeus (Carl von Linné) designed

14616-401: The toes are often webbed for swimming or have suction pads for climbing. Frogs have large eyes and no tail. Salamanders resemble lizards in appearance; their short legs project sideways, the belly is close to or in contact with the ground and they have a long tail. Caecilians superficially resemble earthworms and are limbless. They burrow by means of zones of muscle contractions which move along

14742-555: The transcription factor Brn3b is expressed by M1 ipRGCs that target the OPN, but not by ones that target the SCN. Using this knowledge, they designed an experiment to cross Melanopsin- Cre mice with mice that conditionally expressed a toxin from the Brn3b locus. This allowed them to selectively ablate only the OPN projecting M1 ipRGCS, resulting in a loss of pupil reflexes. However, this did not impair circadian photo entrainment. This demonstrated that

14868-538: The trunk, which is held well clear of the ground. The bones of mammals are well ossified and their teeth, which are usually differentiated, are coated in a layer of prismatic enamel . The teeth are shed once ( milk teeth ) during the animal's lifetime or not at all, as is the case in cetaceans . Mammals have three bones in the middle ear and a cochlea in the inner ear . They are clothed in hair and their skin contains glands which secrete sweat . Some of these glands are specialized as mammary glands , producing milk to feed

14994-442: The vertebrae. Their skulls are mostly broad and short, and are often incompletely ossified. Their skin contains little keratin and lacks scales, but contains many mucous glands and in some species, poison glands. The hearts of amphibians have three chambers, two atria and one ventricle . They have a urinary bladder and nitrogenous waste products are excreted primarily as urea . Amphibians breathe by means of buccal pumping ,

15120-437: The water column. Amphibians are a class of animals comprising frogs , salamanders and caecilians . They are tetrapods , but the caecilians and a few species of salamander have either no limbs or their limbs are much reduced in size. Their main bones are hollow and lightweight and are fully ossified and the vertebrae interlock with each other and have articular processes . Their ribs are usually short and may be fused to

15246-411: The young. Mammals breathe with lungs and have a muscular diaphragm separating the thorax from the abdomen which helps them draw air into the lungs. The mammalian heart has four chambers, and oxygenated and deoxygenated blood are kept entirely separate. Nitrogenous waste is excreted primarily as urea. Mammals are amniotes , and most are viviparous , giving birth to live young. Exceptions to this are

15372-461: Was a direct relationship between irregular light exposure, aberrant mood, cognitive function, normal sleep patterns and circadian oscillations. In a study published in 2012, the Hattar Laboratory was able to show that deviant light cycles directly induce depression -like symptoms and lead to impaired learning in mice, independent of sleep and circadian oscillations. ipRGCs project to areas of

15498-564: Was also affected, with T7 mice failing to show preference for novel objects in the novel object recognition test. Mice without (Opn4 mice) are not susceptible to the negative effects of an aberrant light cycle, indicating that light information transmitted through these cells plays an important role in regulation of mood and cognitive functions such as learning and memory. More recently, light therapy and melatonin administration have been explored by Alfred J. Lewy ( OHSU ), Josephine Arendt ( University of Surrey , UK) and other researchers as

15624-425: Was also responsible for naming and describing the function of the epiglottis and the heart's valves, including the tricuspid. During the third century, Greek physicians were able to differentiate nerves from blood vessels and tendons and to realize that the nerves convey neural impulses. It was Herophilus who made the point that damage to motor nerves induced paralysis. Herophilus named the meninges and ventricles in

15750-421: Was particularly concerned with studies of the circulatory and nervous systems. He could distinguish the human body's sensory and motor nerves and believed air entered the lungs and heart, which was then carried throughout the body. His distinction between the arteries and veins—the arteries carrying the air through the body, while the veins carry the blood from the heart was a great anatomical discovery. Erasistratus

15876-470: Was the standard anatomy textbook for the next century. Pupillary light reflex The pupillary light reflex ( PLR ) or photopupillary reflex is a reflex that controls the diameter of the pupil , in response to the intensity ( luminance ) of light that falls on the retinal ganglion cells of the retina in the back of the eye , thereby assisting in adaptation of vision to various levels of lightness/darkness. A greater intensity of light causes

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