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Saturday, December 11, 2021

Human body

From Wikipedia, the free encyclopedia
Female (left) and male (right) adult human bodies photographed in ventral (above) and dorsal (below) perspectives. Naturally-occurring pubic, body, and facial hair has been deliberately removed to show anatomy

The human body is the structure of a human being. It is composed of many different types of cells that together create tissues and subsequently organ systems. They ensure homeostasis and the viability of the human body.

It comprises a head, neck, trunk (which includes the thorax and abdomen), arms and hands, legs and feet.

The study of the human body involves anatomy, physiology, histology and embryology. The body varies anatomically in known ways. Physiology focuses on the systems and organs of the human body and their functions. Many systems and mechanisms interact in order to maintain homeostasis, with safe levels of substances such as sugar and oxygen in the blood.

The body is studied by health professionals, physiologists, anatomists, and by artists to assist them in their work.

Composition

Elements of the human body by mass. Trace elements are less than 1% combined (and each less than 0.1%).
201 Elements of the Human Body.02.svg Element Symbol Percent mass Percent atoms
Oxygen O 65.0 24.0
Carbon C 18.5 12.0
Hydrogen H 9.5 62.0
Nitrogen N 3.2 1.1
Calcium Ca 1.5 0.22
Phosphorus P 1.0 0.22
Potassium K 0.4 0.03
Sulfur S 0.3 0.038
Sodium Na 0.2 0.037
Chlorine Cl 0.2 0.024
Magnesium Mg 0.1 0.015
Trace elements
< 0.1 < 0.3

The human body is composed of elements including hydrogen, oxygen, carbon, calcium and phosphorus. These elements reside in trillions of cells and non-cellular components of the body.

The adult male body is about 60% water for a total water content of some 42 litres (9.2 imp gal; 11 US gal). This is made up of about 19 litres (4.2 imp gal; 5.0 US gal) of extracellular fluid including about 3.2 litres (0.70 imp gal; 0.85 US gal) of blood plasma and about 8.4 litres (1.8 imp gal; 2.2 US gal) of interstitial fluid, and about 23 litres (5.1 imp gal; 6.1 US gal) of fluid inside cells. The content, acidity and composition of the water inside and outside cells is carefully maintained. The main electrolytes in body water outside cells are sodium and chloride, whereas within cells it is potassium and other phosphates.

Cells

The body contains trillions of cells, the fundamental unit of life. At maturity, there are roughly 30–37 trillion cells in the body, an estimate arrived at by totaling the cell numbers of all the organs of the body and cell types. The body is also host to about the same number of non-human cells as well as multicellular organisms which reside in the gastrointestinal tract and on the skin. Not all parts of the body are made from cells. Cells sit in an extracellular matrix that consists of proteins such as collagen, surrounded by extracellular fluids. Of the 70 kg (150 lb) weight of an average human body, nearly 25 kg (55 lb) is non-human cells or non-cellular material such as bone and connective tissue.

Genome

Cells in the body function because of DNA. DNA sits within the nucleus of a cell. Here, parts of DNA are copied and sent to the body of the cell via RNA. The RNA is then used to create proteins which form the basis for cells, their activity, and their products. Proteins dictate cell function and gene expression, a cell is able to self-regulate by the amount of proteins produced. However, not all cells have DNA; some cells such as mature red blood cells lose their nucleus as they mature.

Tissues

External video
2120 Major Systemic Artery.jpg
video icon Human Body 101, National Geographic, 5:10

The body consists of many different types of tissue, defined as cells that act with a specialised function. The study of tissues is called histology and often occurs with a microscope. The body consists of four main types of tissues. These are lining cells (epithelia), connective tissue, nerve tissue and muscle tissue.

Cells that lie on surfaces exposed to the outside world or gastrointestinal tract (epithelia) or internal cavities (endothelium) come in numerous shapes and forms – from single layers of flat cells, to cells with small beating hair-like cilia in the lungs, to column-like cells that line the stomach. Endothelial cells are cells that line internal cavities including blood vessels and glands. Lining cells regulate what can and can't pass through them, protect internal structures, and function as sensory surfaces.

Organs

Organs, structured collections of cells with a specific function, mostly sit within the body, with the exception of skin. Examples include the heart, lungs and liver. Many organs reside within cavities within the body. These cavities include the abdomen (which contains the stomach, for example) and pleura, which contains the lungs.

Heart

The heart is an organ located in the thoracic cavity between the lungs and slightly to the left. It is surrounded by the pericardium which holds it in place in the mediastinum and serves to protect it from blunt trauma, infection and help lubricate the movement of the heart via pericardial fluid. The heart works by pumping blood around the body allowing oxygen, nutrients, waste, hormones and white blood cells to be transported.

Diagram of the human heart

The heart is composed of two atrium and two ventricles. The primary purpose of the atrium is to allow uninterrupted venous blood flow to the heart during ventricular systole. This allows enough blood to get into the ventricles during atrial systole. A lack of the atrium would cause a decrease in cardiac output of 75%. The purpose of the ventricles is to pump blood to the lungs through the right ventricle and to the rest of the body through the left ventricle.

The heart has an electrical conduction system to control the contractions and relaxation of the muscle. It starts in the sinoatrial node traveling through the atria causing them to pump blood into the ventricles. It then travels to the atrioventricular node which makes the signal slow down slightly allowing the ventricles to fill with blood before pumping it out and starting the cycle over again.

Coronary artery disease is the leading cause of death worldwide, making up 16% of all deaths. It is caused by the buildup of plaque in the coronary arteries supplying the heart, eventually the arteries may become so narrow that not enough blood is able to reach the myocardium, a condition known as myocardial infarction or heart attack, this can cause heart failure or cardiac arrest and eventually death. Risk factors for coronary artery disease include obesity, smoking, high cholesterol, high blood pressure, lack of exercise and diabetes. Cancer can affect the heart, though it is exceedingly rare and has usually metastasized from another part of the body such as the lungs or breasts. This is because the heart cells quickly stop dividing and all growth occurs through size increase rather than cell division.

Gallbladder

The gallbladder is a hollow pear shaped organ located posterior to the inferior middle part of the right lobe of the liver. It is variable in shape and size. It stores bile before it is released into the small intestine via the common bile duct to help with digestion of fats. It receives bile from the liver via the cystic duct which connects to the common hepatic duct to form the common bile duct.

The gallbladder gets its blood supply from the cystic artery which in most people emerges from the right hepatic artery.

Gallstones is a common disease in which one or more stones form in the gallbladder or biliary tract. Most people are asymptomatic but if a stone blocks the biliary tract it causes a gallbladder attack, symptoms may include sudden pain in the upper right abdomen and or center of the abdomen. Nausea and vomiting may also occur. Typical treatment is removal of the gallbladder through a procedure called a cholecystectomy. Having gallstones is a risk factor for gallbladder cancer which although quite uncommon is rapidly fatal if not diagnosed early.

Systems

Circulatory system

The circulatory system consists of the heart and blood vessels (arteries, veins and capillaries). The heart propels the circulation of the blood, which serves as a "transportation system" to transfer oxygen, fuel, nutrients, waste products, immune cells and signalling molecules (i.e. hormones) from one part of the body to another. Paths of blood circulation within the human body can be divided into two circuits: the pulmonary circuit, which pumps blood to the lungs to receive oxygen and leave carbon dioxide, and the systemic circuit, which carries blood from the heart off to the rest of the body. The blood consists of fluid that carries cells in the circulation, including some that move from tissue to blood vessels and back, as well as the spleen and bone marrow.

Digestive system

Digestive system

The digestive system consists of the mouth including the tongue and teeth, esophagus, stomach, (gastrointestinal tract, small and large intestines, and rectum), as well as the liver, pancreas, gallbladder, and salivary glands. It converts food into small, nutritional, non-toxic molecules for distribution and absorption into the body. These molecules take the form of proteins (which are broken down into amino acids), fats, vitamins and minerals (the last of which are mainly ionic rather than molecular). After being swallowed, food moves through the gastrointestinal tract by means of peristalsis: the systematic expansion and contraction of muscles to push food from one area to the next.

Digestion begins in the mouth, which chews food into smaller pieces for easier digestion. Then it is swallowed, and moves through the esophagus to the stomach. In the stomach, food is mixed with gastric acids to allow the extraction of nutrients. What is left is called chyme; this then moves into the small intestine, which absorbs the nutrients and water from the chyme. What remains passes on to the large intestine, where it is dried to form feces; these are then stored in the rectum until they are expelled through the anus.

Endocrine system

The endocrine system consists of the principal endocrine glands: the pituitary, thyroid, adrenals, pancreas, parathyroids, and gonads, but nearly all organs and tissues produce specific endocrine hormones as well. The endocrine hormones serve as signals from one body system to another regarding an enormous array of conditions, and resulting in variety of changes of function.

Immune system

The immune system consists of the white blood cells, the thymus, lymph nodes and lymph channels, which are also part of the lymphatic system. The immune system provides a mechanism for the body to distinguish its own cells and tissues from outside cells and substances and to neutralize or destroy the latter by using specialized proteins such as antibodies, cytokines, and toll-like receptors, among many others.

Skin

Integumentary system

The integumentary system consists of the covering of the body (the skin), including hair and nails as well as other functionally important structures such as the sweat glands and sebaceous glands. The skin provides containment, structure, and protection for other organs, and serves as a major sensory interface with the outside world.

Lymphatic system

The lymphatic system extracts, transports and metabolizes lymph, the fluid found in between cells. The lymphatic system is similar to the circulatory system in terms of both its structure and its most basic function, to carry a body fluid.

Musculoskeletal system

The musculoskeletal system consists of the human skeleton (which includes bones, ligaments, tendons, and cartilage) and attached muscles. It gives the body basic structure and the ability for movement. In addition to their structural role, the larger bones in the body contain bone marrow, the site of production of blood cells. Also, all bones are major storage sites for calcium and phosphate. This system can be split up into the muscular system and the skeletal system.

Nervous system

Nervous system

The nervous system consists of the body's neurons and glial cells, which together form the nerves, ganglia and gray matter which in turn form the brain and related structures. The brain is the organ of thought, emotion, memory, and sensory processing; it serves many aspects of communication and controls various systems and functions. The special senses consist of vision, hearing, taste, and smell. The eyes, ears, tongue, and nose gather information about the body's environment.

From a structural perspective, the nervous system is typically subdivided into two component parts: the central nervous system (CNS), composed of the brain and the spinal cord; and the peripheral nervous system (PNS), composed of the nerves and ganglia outside the brain and spinal cord. The CNS is mostly responsible for organizing motion, processing sensory information, thought, memory, cognition and other such functions. It remains a matter of some debate whether the CNS directly gives rise to consciousness. The peripheral nervous system (PNS) is mostly responsible for gathering information with sensory neurons and directing body movements with motor neurons.

From a functional perspective, the nervous system is again typically divided into two component parts: the somatic nervous system (SNS) and the autonomic nervous system (ANS). The SNS is involved in voluntary functions like speaking and sensory processes. The ANS is involved in involuntary processes, such as digestion and regulating blood pressure.

The nervous system is subject to many different diseases. In epilepsy, abnormal electrical activity in the brain can cause seizures. In multiple sclerosis, the immune system attacks the nerve linings, damaging the nerves' ability to transmit signals. Amyotrophic lateral sclerosis (ALS), also known as Lou Gehrig's disease, is a motor neuron disease which gradually reduces movement in patients. There are also many other diseases of the nervous system.

Reproductive system

The primary purpose of the reproductive system is to produce new humans in the form of children and ensure their sexual development so that they can produce new humans too. It is composed of the gonads (testes and ovaries) which produce eggs and sperm cells (gametes) and produce hormones necessary for proper sexual development. The rest of the glands and ducts of the reproductive system are responsible for the transportation and sustaining of the gametes and to nurture the offspring.

The reproductive system is first stimulated by the hypothalamus during puberty which causes the ovaries and testes to produce testosterone (male) and estrogen and progesterone (female). Male puberty generally occurs between the ages of 13-15 and is characterized by beginning of sperm production and the development of secondary sex characteristics such as increased height and weight, broadened shoulders, pubic and facial hair, voice deepening and muscle development. Female puberty generally occurs between the ages of 9-13 and is characterized by ovulation and menstruation. The growth of secondary sex characteristics such as growth of pubic and underarm hair, breast, uterine and vaginal growth, widening hips and increased height and weight also occur during the female puberty process.

The external male reproductive system is made up of the penis and the scrotum which is a bag that protects the testes. The penis consists of the glans which is the head of the penis and contains the urethra and urinary meatus the point where urine exits the penis, the rest of the penis is called the shaft or corpus cavernosum and contains most of the urethra. The glans is covered by a fold of skin called the foreskin though this can be removed through circumcision.

External female reproductive system with mons pubis highlighted

Internally the male reproductive system starts in the testes where hundreds of seminiferous tubules produce sperm which is then stored and matured into spermatozoon in the epididymis. They are then brought through the vas deferens which leads away from the testes to the seminal vesicles where the sperm cells are mixed with a fructose-rich fluid from the seminal vesicles allowing the sperm to stay alive and remain healthy. It is then brought by the ejaculatory duct through the prostate and bulbourethral gland, and mixed with fluids from said glands. The fluid from the prostate helps to neutralize the acidity of the vagina and keep the sperm alive. This mixture of sperm and accessory gland fluids is called semen and is released through the urethra during ejaculation. Almost the entire internal male reproductive system works in pairs with two testes, two epididymis, two vas deferens, two seminal vesicles, two ejaculatory ducts, two bulbourethral glands, but only one prostate and urethra.

The external female reproductive system also called the vulva consists of the mons pubis a fatty mass that covers the pubis, the labia majora (outer lips of the vagina), the labia minora (inner lips of the vagina), the vaginal opening which opens into vagina and is where the penis is inserted during sexual intercourse and children are born, the urethral opening which is the opening for the urethra that carries urine from the bladder and the clitoris which contains the most sensitive nerve endings and in humans has no use other than sexual pleasure. Located between the anus and vagina is the perineum.

Internally the female reproductive system contains two ovaries, the uterus, two fallopian tubes and the cervix. At birth a female has about 700,000 oocytes (the immature version of the egg cell) in both ovaries combined, though this degenerates to about 400,000 by the time puberty is reached. This is a lifetime supply as after birth no more oocytes are produced, compared to males where sperm cells are produced during their entire lifetime. During puberty the menstrual cycle begins for the first time, in response to low estrogen and progesterone levels the hypothalamus releases gonadotropin releasing hormone (GnRH). This causes the anterior pituitary gland to release follicle-stimulating hormone (FSH) and luteinizing hormone (LH). FSH stimulates ovarian follicles to grow and one dominant follicle will eventually take over. As it continues to grow it releases more and more estradiol which stimulates LH secretion and suppresses FSH secretion preventing further follicular growth. When LH levels are highest the follicle ruptures releasing the ovum in a process called ovulation where it is then moved to one of the fallopian tubes. After ovulation the portion of the follicle that remains in the ovary is transformed into corpus luteum which continues to produce estrogen and high levels of progesterone. The progesterone causes the endometrium to grow thick preparing it for implantation of a fertilized egg. If fertilization occurs the corpus luteum continues to secrete hormones until the placenta develops enough to secrete the necessary hormones for maintaining pregnancy. Eventually the corpus luteum will turn into corpus albicans which is essentially scar tissue. If fertilization fails the corpus luteum will degrade into corpus albicans and stop secreting enough progesterone and estrogen causing the endometrial lining to break resulting in menstruation.

Many diseases affect the reproductive system such as polycystic ovary syndrome (PCOS) which is characterized by elevated androgen levels, menstrual irregularities and/or small cysts on one or both of the ovaries. It is a common disease affecting at least 7% of adult women. Symptoms include excess body hair, infertility, weight gain, male pattern baldness and irregular menstruation.

Another reproductive disease is testicular torsion, it occurs when the spermatic cord which holds up the testicle wraps around itself cutting off blood flow to the testicle. Testicular torsion is a medical emergency and can result in testicular death or infertility if not treated immediately by physically unwrapping the spermatic cord. If the testicle has suffered severe damage, it may need to be surgically in a lateral orchiectomy. The primary symptom of testicular torsion is severe testicular pain at rest, nausea and vomiting may also occur. Testicular torsion can occur at any age though it is most common in 12-18 year olds.

Sexually transmitted infections such as syphilis, HIV, chlamydia, HPV and genital warts are spread through sexual intercourse including oral, vaginal and anal sex. Many of these infections can be lethal if left untreated though others are mostly harmless.

Cancer can affect many parts of the reproductive system including the penis, testicles, prostate, ovaries, cervix, vagina, fallopian, uterus and vulva.

Respiratory system

The respiratory system consists of the nose, nasopharynx, trachea, and lungs. It brings oxygen from the air and excretes carbon dioxide and water back into the air. First, air is pulled through the trachea into the lungs by the diaphragm pushing down, which creates a vacuum. Air is briefly stored inside small sacs known as alveoli (sing.: alveolus) before being expelled from the lungs when the diaphragm contracts again. Each alveolus is surrounded by capillaries carrying deoxygenated blood, which absorbs oxygen out of the air and into the bloodstream.

For the respiratory system to function properly, there need to be as few impediments as possible to the movement of air within the lungs. Inflammation of the lungs and excess mucus are common sources of breathing difficulties. In asthma, the respiratory system is persistently inflamed, causing wheezing and/or shortness of breath. Pneumonia occurs through infection of the alveoli, and may be caused by tuberculosis. Emphysema, commonly a result of smoking, is caused by damage to connections between the alveoli.

Urinary system

Female urinary system

The urinary system consists of the two kidneys, two ureters, bladder, and urethra. It removes waste materials from the blood through urine, which carries a variety of waste molecules and excess ions and water out of the body.

First, the kidneys filter the blood through their respective nephrons, removing waste products like urea, creatinine and maintaining the proper balance of electrolytes and turning the waste products into urine by combining them with water from the blood. The kidneys filter about 150 quarts (170 liters) of blood daily, but most of it is returned to the blood stream with only 1-2 quarts (1-2 liters) ending up as urine. The urine is brought by the ureters from the kidneys down to the bladder.

The smooth muscle lining the ureter walls continuously tighten and relax through a process called peristalsis to force urine away from the kidneys and down into the bladder. Small amounts of urine are released into the bladder every 10-15 seconds.

The bladder is a hollow balloon shaped organ located in the pelvis. It stores urine until the brain signals it to relax the urinary sphincter and release the urine into the urethra starting urination. A normal bladder can hold up to 16 ounces (half a liter) for 3-5 hours comfortably.

Numerous diseases affect the urinary system including kidney stones which are formed when materials in the urine concentrate enough to form a solid mass, urinary tract infections which are infections of the urinary tract and can cause pain when urinating, frequent urination and even death if left untreated. Renal failure occurs when the kidneys fail to adequately filter waste from the blood and can lead to death if not treated with dialysis or kidney transplantation. Cancer can affect the bladder, kidneys, urethra and ureters with the latter two being far more rare.

Anatomy

Cavities of human body

Human anatomy is the study of the shape and form of the human body. The human body has four limbs (two arms and two legs), a head and a neck which connect to the torso. The body's shape is determined by a strong skeleton made of bone and cartilage, surrounded by fat, muscle, connective tissue, organs, and other structures. The spine at the back of the skeleton contains the flexible vertebral column which surrounds the spinal cord, which is a collection of nerve fibres connecting the brain to the rest of the body. Nerves connect the spinal cord and brain to the rest of the body. All major bones, muscles, and nerves in the body are named, with the exception of anatomical variations such as sesamoid bones and accessory muscles.

Blood vessels carry blood throughout the body, which moves because of the beating of the heart. Venules and veins collect blood low in oxygen from tissues throughout the body. These collect in progressively larger veins until they reach the body's two largest veins, the superior and inferior vena cava, which drain blood into the right side of the heart. From here, the blood is pumped into the lungs where it receives oxygen and drains back into the left side of the heart. From here, it is pumped into the body's largest artery, the aorta, and then progressively smaller arteries and arterioles until it reaches tissue. Here blood passes from small arteries into capillaries, then small veins and the process begins again. Blood carries oxygen, waste products, and hormones from one place in the body to another. Blood is filtered at the kidneys and liver.

The body consists of a number of body cavities, separated areas which house different organ systems. The brain and central nervous system reside in an area protected from the rest of the body by the blood brain barrier. The lungs sit in the pleural cavity. The intestines, liver, and spleen sit in the abdominal cavity.

Height, weight, shape and other body proportions vary individually and with age and sex. Body shape is influenced by the distribution of bones, muscle and fat tissue.

Physiology

Human physiology is the study of how the human body functions. This includes the mechanical, physical, bioelectrical, and biochemical functions of humans in good health, from organs to the cells of which they are composed. The human body consists of many interacting systems of organs. These interact to maintain homeostasis, keeping the body in a stable state with safe levels of substances such as sugar and oxygen in the blood.

Each system contributes to homeostasis, of itself, other systems, and the entire body. Some combined systems are referred to by joint names. For example, the nervous system and the endocrine system operate together as the neuroendocrine system. The nervous system receives information from the body, and transmits this to the brain via nerve impulses and neurotransmitters. At the same time, the endocrine system releases hormones, such as to help regulate blood pressure and volume. Together, these systems regulate the internal environment of the body, maintaining blood flow, posture, energy supply, temperature, and acid balance (pH).

Development

Baby being carried

Development of the human body is the process of growth to maturity. The process begins with fertilisation, where an egg released from the ovary of a female is penetrated by sperm. The egg then lodges in the uterus, where an embryo and later fetus develop until birth. Growth and development occur after birth, and include both physical and psychological development, influenced by genetic, hormonal, environmental and other factors. Development and growth continue throughout life, through childhood, adolescence, and through adulthood to old age, and are referred to as the process of aging.

Society and culture

Professional study

Anatomical study by Leonardo da Vinci

Health professionals learn about the human body from illustrations, models, and demonstrations. Medical and dental students in addition gain practical experience, for example by dissection of cadavers. Human anatomy, physiology, and biochemistry are basic medical sciences, generally taught to medical students in their first year at medical school.

Depiction

Figure drawing by Lovis Corinth (before 1925)

Anatomy has served the visual arts since Ancient Greek times, when the 5th century BC sculptor Polykleitos wrote his Canon on the ideal proportions of the male nude. In the Italian Renaissance, artists from Piero della Francesca (c. 1415–1492) onwards, including Leonardo da Vinci (1452–1519) and his collaborator Luca Pacioli (c. 1447–1517), learnt and wrote about the rules of art, including visual perspective and the proportions of the human body.

History of anatomy

Two facing pages of text with woodcuts of naked male and female figures, in the Epitome by Andreas Vesalius, 1543

In Ancient Greece, the Hippocratic Corpus described the anatomy of the skeleton and muscles. The 2nd century physician Galen of Pergamum compiled classical knowledge of anatomy into a text that was used throughout the Middle Ages. In the Renaissance, Andreas Vesalius (1514–1564) pioneered the modern study of human anatomy by dissection, writing the influential book De humani corporis fabrica. Anatomy advanced further with the invention of the microscope and the study of the cellular structure of tissues and organs. Modern anatomy uses techniques such as magnetic resonance imaging, computed tomography, fluoroscopy and ultrasound imaging to study the body in unprecedented detail.

History of physiology

The study of human physiology began with Hippocrates in Ancient Greece, around 420 BCE, and with Aristotle (384–322 BCE) who applied critical thinking and emphasis on the relationship between structure and function. Galen (ca. 126–199) was the first to use experiments to probe the body's functions. The term physiology was introduced by the French physician Jean Fernel (1497–1558). In the 17th century, William Harvey (1578–1657) described the circulatory system, pioneering the combination of close observation with careful experiment. In the 19th century, physiological knowledge began to accumulate at a rapid rate with the cell theory of Matthias Schleiden and Theodor Schwann in 1838, that organisms are made up of cells. Claude Bernard (1813–1878) created the concept of the milieu interieur (internal environment), which Walter Cannon (1871–1945) later said was regulated to a steady state in homeostasis. In the 20th century, the physiologists Knut Schmidt-Nielsen and George Bartholomew extended their studies to comparative physiology and ecophysiology. Most recently, evolutionary physiology has become a distinct subdiscipline.

Fertilisation

From Wikipedia, the free encyclopedia
 
Sperm and ovum fusing

Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a new individual organism or offspring and initiate its development. Processes such as insemination or pollination which happen before the fusion of gametes are also sometimes informally called fertilisation. The cycle of fertilisation and development of new individuals is called sexual reproduction. During double fertilisation in angiosperms the haploid male gamete combines with two haploid polar nuclei to form a triploid primary endosperm nucleus by the process of vegetative fertilisation.

History

In Antiquity, Aristotle conceived the formation of new individuals through fusion of male and female fluids, with form and function emerging gradually, in a mode called by him as epigenetic.

In 1784, Spallanzani established the need of interaction between the female's ovum and male's sperm to form a zygote in frogs. In 1827, von Baer observed a therian mammalian egg for the first time. Oscar Hertwig (1876), in Germany, described the fusion of nuclei of spermatozoa and of ova from sea urchin.

Evolution

The evolution of fertilisation is related to the origin of meiosis, as both are part of sexual reproduction, originated in eukaryotes. There are two conflicting theories on how the couple meiosis–fertilisation arose. One is that it evolved from prokaryotic sex (bacterial recombination) as eukaryotes evolved from prokaryotes. The other is that mitosis originated meiosis.

Fertilisation in plants

In the Bryophyte land plants, fertilisation takes place within the archegonium. This moss has been genetically modified so that the unfertilised egg within the archegonium produces a blue colour.

The gametes that participate in fertilisation of plants are the sperm (male) and the egg (female) cell. Various families of plants have differing methods by which the gametes produced by the male and female gametophytes come together and are fertilised. In Bryophyte land plants, fertilisation of the sperm and egg takes place within the archegonium. In seed plants, the male gametophyte is called a pollen grain. After pollination, the pollen grain germinates, and a pollen tube grows and penetrates the ovule through a tiny pore called a micropyle. The sperm are transferred from the pollen through the pollen tube to the ovule where the egg is fertilised. In flowering plants, two sperm cells are released from the pollen grain, and a second fertilisation event involving the second sperm cell and the central cell of the ovule, which is a second female gamete.

Pollen tube growth

Unlike animal sperm which is motile, plant sperm is immotile and relies on the pollen tube to carry it to the ovule where the sperm is released. The pollen tube penetrates the stigma and elongates through the extracellular matrix of the style before reaching the ovary. Then near the receptacle, it breaks through the ovule through the micropyle (an opening in the ovule wall) and the pollen tube "bursts" into the embryo sac, releasing sperm. The growth of the pollen tube has been believed to depend on chemical cues from the pistil, however these mechanisms were poorly understood until 1995. Work done on tobacco plants revealed a family of glycoproteins called TTS proteins that enhanced growth of pollen tubes. Pollen tubes in a sugar free pollen germination medium and a medium with purified TTS proteins both grew. However, in the TTS medium, the tubes grew at a rate 3x that of the sugar-free medium. TTS proteins were also placed on various locations of semi in vevo pollinated pistils, and pollen tubes were observed to immediately extend toward the proteins. Transgenic plants lacking the ability to produce TTS proteins exhibited slower pollen tube growth and reduced fertility.

Rupture of pollen tube

The rupture of the pollen tube to release sperm in Arabidopsis has been shown to depend on a signal from the female gametophyte. Specific proteins called FER protein kinases present in the ovule control the production of highly reactive derivatives of oxygen called reactive oxygen species (ROS). ROS levels have been shown via GFP to be at their highest during floral stages when the ovule is the most receptive to pollen tubes, and lowest during times of development and following fertilisation. High amounts of ROS activate Calcium ion channels in the pollen tube, causing these channels to take up Calcium ions in large amounts. This increased uptake of calcium causes the pollen tube to rupture, and release its sperm into the ovule. Pistil feeding assays in which plants were fed diphenyl iodonium chloride (DPI) suppressed ROS concentrations in Arabidopsis, which in turn prevented pollen tube rupture.

Bryophytes

Bryophyte is a traditional name used to refer to all embryophytes (land plants) that do not have true vascular tissue and are therefore called "non-vascular plants". Some bryophytes do have specialised tissues for the transport of water; however, since these do not contain lignin, they are not considered true vascular tissue.

Ferns

A fern is a member of a group of roughly 12,000 species of vascular plants that reproduce via spores and have neither seeds nor flowers. They differ from mosses by being vascular (i.e. having water-conducting vessels). They have stems and leaves, like other vascular plants. Most ferns have what are called fiddleheads that expand into fronds, which are each delicately divided.

Gymnosperms

The gymnosperms are a group of seed producing plants that includes conifers, Cycads, Ginkgo, and Gnetales. The term "gymnosperm" comes from the Greek composite word γυμνόσπερμος (γυμνός gymnos, "naked" and σπέρμα sperma, "seed"), meaning "naked seeds", after the unenclosed condition of their seeds (called ovules in their unfertilised state). Their naked condition stands in contrast to the seeds and ovules of flowering plants (angiosperms), which are enclosed within an ovary. Gymnosperm seeds develop either on the surface of scales or leaves, often modified to form cones, or at the end of short stalks as in Ginkgo.

Flowering plants

After being fertilised, the ovary starts to swell and develop into the fruit. With multi-seeded fruits, multiple grains of pollen are necessary for syngamy with each ovule. The growth of the pollen tube is controlled by the vegetative (or tube) cytoplasm. Hydrolytic enzymes are secreted by the pollen tube that digest the female tissue as the tube grows down the stigma and style; the digested tissue is used as a nutrient source for the pollen tube as it grows. During pollen tube growth towards the ovary, the generative nucleus divides to produce two separate sperm nuclei (haploid number of chromosomes) – a growing pollen tube therefore contains three separate nuclei, two sperm and one tube. The sperms are interconnected and dimorphic, the large one, in a number of plants, is also linked to the tube nucleus and the interconnected sperm and the tube nucleus form the "male germ unit".

Double fertilisation is the process in angiosperms (flowering plants) in which two sperm from each pollen tube fertilise two cells in a female gametophyte (sometimes called an embryo sac) that is inside an ovule. After the pollen tube enters the gametophyte, the pollen tube nucleus disintegrates and the two sperm cells are released; one of the two sperm cells fertilises the egg cell (at the bottom of the gametophyte near the micropyle), forming a diploid (2n) zygote. This is the point when fertilisation actually occurs; pollination and fertilisation are two separate processes. The nucleus of the other sperm cell fuses with two haploid polar nuclei (contained in the central cell) in the centre of the gametophyte. The resulting cell is triploid (3n). This triploid cell divides through mitosis and forms the endosperm, a nutrient-rich tissue, inside the seed.

The two central-cell maternal nuclei (polar nuclei) that contribute to the endosperm arise by mitosis from the single meiotic product that also gave rise to the egg. Therefore, maternal contribution to the genetic constitution of the triploid endosperm is double that of the embryo.

One primitive species of flowering plant, Nuphar polysepala, has endosperm that is diploid, resulting from the fusion of a sperm with one, rather than two, maternal nuclei. It is believed that early in the development of angiosperm linages, there was a duplication in this mode of reproduction, producing seven-celled/eight-nucleate female gametophytes, and triploid endosperms with a 2:1 maternal to paternal genome ratio.

In many plants, the development of the flesh of the fruit is proportional to the percentage of fertilised ovules. For example, with watermelon, about a thousand grains of pollen must be delivered and spread evenly on the three lobes of the stigma to make a normal sized and shaped fruit.

Cross-fertilisation and self-fertilisation represent different strategies with differing benefits and costs. An estimated 48.7% of plant species are either dioecious or self-incompatible obligate out-crossers. It is also estimated that about 42% of flowering plants exhibit a mixed mating system in nature.

In the most common kind of mixed mating system, individual plants produce a single type of flower and fruits may contain self-fertilised, out-crossed or a mixture of progeny types. The transition from cross-fertilisation to self-fertilisation is the most common evolutionary transition in plants, and has occurred repeatedly in many independent lineages. About 10-15% of flowering plants are predominantly self-fertilising.

Self-pollination

Under circumstances where pollinators or mates are rare, self-fertilisation offers the advantage of reproductive assurance. Self-fertilisation can therefore result in improved colonisation ability. In some species, self-fertilisation has persisted over many generations. Capsella rubella is a self-fertilisating species that became self-compatible 50,000 to 100,000 years ago. Arabidopsis thaliana is a predominantly self-fertilising plant with an out-crossing rate in the wild of less than 0.3%; a study suggested that self-fertilisation evolved roughly a million years ago or more in A. thaliana. In long-established self-fertilising plants, the masking of deleterious mutations and the production of genetic variability is infrequent and thus unlikely to provide a sufficient benefit over many generations to maintain the meiotic apparatus. Consequently, one might expect self-fertilisation to be replaced in nature by an ameiotic asexual form of reproduction that would be less costly. However the actual persistence of meiosis and self-fertilisation as a form of reproduction in long-established self-fertilising plants may be related to the immediate benefit of efficient recombinational repair of DNA damage during formation of germ cells provided by meiosis at each generation.

Fertilisation in animals

The mechanics behind fertilisation has been studied extensively in sea urchins and mice. This research addresses the question of how the sperm and the appropriate egg find each other and the question of how only one sperm gets into the egg and delivers its contents. There are three steps to fertilisation that ensure species-specificity:

  1. Chemotaxis
  2. Sperm activation/acrosomal reaction
  3. Sperm/egg adhesion

Internal vs. external

Consideration as to whether an animal (more specifically a vertebrate) uses internal or external fertilisation is often dependent on the method of birth. Oviparous animals laying eggs with thick calcium shells, such as chickens, or thick leathery shells generally reproduce via internal fertilisation so that the sperm fertilises the egg without having to pass through the thick, protective, tertiary layer of the egg. Ovoviviparous and viviparous animals also use internal fertilisation. It is important to note that although some organisms reproduce via amplexus, they may still use internal fertilisation, as with some salamanders. Advantages to internal fertilisation include: minimal waste of gametes; greater chance of individual egg fertilisation, relatively "longer" time period of egg protection, and selective fertilisation; many females have the ability to store sperm for extended periods of time and can fertilise their eggs at their own desire.

Oviparous animals producing eggs with thin tertiary membranes or no membranes at all, on the other hand, use external fertilisation methods. Such animals may be more precisely termed ovuliparous. Advantages to external fertilisation include: minimal contact and transmission of bodily fluids; decreasing the risk of disease transmission, and greater genetic variation (especially during broadcast spawning external fertilisation methods).

Sea urchins

Acrosome reaction on a sea urchin cell.

Sperm find the eggs via chemotaxis, a type of ligand/receptor interaction. Resact is a 14 amino acid peptide purified from the jelly coat of A. punctulata that attracts the migration of sperm.

After finding the egg, the sperm penetrates the jelly coat through a process called sperm activation. In another ligand/receptor interaction, an oligosaccharide component of the egg binds and activates a receptor on the sperm and causes the acrosomal reaction. The acrosomal vesicles of the sperm fuse with the plasma membrane and are released. In this process, molecules bound to the acrosomal vesicle membrane, such as bindin, are exposed on the surface of the sperm. These contents digest the jelly coat and eventually the vitelline membrane. In addition to the release of acrosomal vesicles, there is explosive polymerisation of actin to form a thin spike at the head of the sperm called the acrosomal process.

The sperm binds to the egg through another ligand reaction between receptors on the vitelline membrane. The sperm surface protein bindin, binds to a receptor on the vitelline membrane identified as EBR1.

Fusion of the plasma membranes of the sperm and egg are likely mediated by bindin. At the site of contact, fusion causes the formation of a fertilisation cone.

Mammals

Mammals internally fertilise through copulation. After a male ejaculates, many sperm move to the upper vagina (via contractions from the vagina) through the cervix and across the length of the uterus to meet the ovum. In cases where fertilisation occurs, the female usually ovulates during a period that extends from hours before copulation to a few days after; therefore, in most mammals it is more common for ejaculation to precede ovulation than vice versa.

When sperm are deposited into the anterior vagina, they are not capable of fertilisation (i.e., non-capacitated) and are characterized by slow linear motility patterns. This motility, combined with muscular contractions enables sperm transport towards the uterus and fallopian tubes. There is a pH gradient within the micro-environment of the female reproductive tract such that the pH near the vaginal opening is lower (approximately 5) than the fallopian tubes (approximately 8). The sperm-specific pH-sensitive calcium transport protein called CatSper increases the sperm cell permeability to calcium as it moves further into the reproductive tract. Intracellular calcium influx contributes to sperm capacitation and hyperactivation, causing a more violent and rapid non-linear motility pattern as sperm approach the oocyte. The capacitated spermatozoon and the oocyte meet and interact in the ampulla of the fallopian tube. Rheotaxis, thermotaxis and chemotaxis are known mechanisms that guide sperm towards the egg during the final stage of sperm migration. Spermatozoa respond (see Sperm thermotaxis) to the temperature gradient of ~2 °C between the oviduct and the ampulla, and chemotactic gradients of progesterone have been confirmed as the signal emanating from the cumulus oophorus cells surrounding rabbit and human oocytes. Capacitated and hyperactivated sperm respond to these gradients by changing their behaviour and moving towards the cumulus-oocyte complex. Other chemotactic signals such as formyl Met-Leu-Phe (fMLF) may also guide spermatozoa.

The zona pellucida, a thick layer of extracellular matrix that surrounds the egg and is similar to the role of the vitelline membrane in sea urchins, binds the sperm. Unlike sea urchins, the sperm binds to the egg before the acrosomal reaction. ZP3, a glycoprotein in the zona pellucida, is responsible for egg/sperm adhesion in mice. The receptor galactosyltransferase (GalT) binds to the N-acetylglucosamine residues on the ZP3 and is important for binding with the sperm and activating the acrosome reaction. ZP3 is sufficient though unnecessary for sperm/egg binding. Two additional sperm receptors exist: a 250kD protein that binds to an oviduct secreted protein, and SED1, which independently binds to the zona. After the acrosome reaction, the sperm is believed to remain bound to the zona pellucida through exposed ZP2 receptors. These receptors are unknown in mice but have been identified in guinea pigs.

In mammals, the binding of the spermatozoon to the GalT initiates the acrosome reaction. This process releases the hyaluronidase that digests the matrix of hyaluronic acid in the vestments around the oocyte. Additionally, heparin-like glycosaminoglycans (GAGs) are released near the oocyte that promote the acrosome reaction. Fusion between the oocyte plasma membranes and sperm follows and allows the sperm nucleus, the typical centriole, and atypical centriole that is attached to the flagellum, but not the mitochondria, to enter the oocyte. The protein CD9 likely mediates this fusion in mice (the binding homolog). The egg "activates" itself upon fusing with a single sperm cell and thereby changes its cell membrane to prevent fusion with other sperm. Zinc atoms are released during this activation.

This process ultimately leads to the formation of a diploid cell called a zygote. The zygote divides to form a blastocyst and, upon entering the uterus, implants in the endometrium, beginning pregnancy. Embryonic implantation not in the uterine wall results in an ectopic pregnancy that can kill the mother.

In such animals as rabbits, coitus induces ovulation by stimulating the release of the pituitary hormone gonadotropin; this release greatly increases the likelihood of pregnancy.

Humans

Fertilisation in humans. The sperm and ovum unite through fertilisation, creating a zygote that (over the course of 8-9 days) implants in the uterine wall, where it resides for nine months.

Fertilisation in humans is the union of a human egg and sperm, usually occurring in the ampulla of the fallopian tube, producing a zygote cell, or fertilised egg, initiating prenatal development. Scientists discovered the dynamics of human fertilisation in the nineteenth century.

The term conception commonly refers to "the process of becoming pregnant involving fertilization or implantation or both". Its use makes it a subject of semantic arguments about the beginning of pregnancy, typically in the context of the abortion debate. Upon gastrulation, which occurs around 16 days after fertilisation, the implanted blastocyst develops three germ layers, the endoderm, the ectoderm and the mesoderm, and the genetic code of the father becomes fully involved in the development of the embryo; later twinning is impossible. Additionally, interspecies hybrids survive only until gastrulation and cannot further develop. However, some human developmental biology literature refers to the conceptus and such medical literature refers to the "products of conception" as the post-implantation embryo and its surrounding membranes. The term "conception" is not usually used in scientific literature because of its variable definition and connotation.

Insects

Red-veined darters (Sympetrum fonscolombii) flying "in cop" (male ahead), enabling the male to prevent other males from mating. The eggs are fertilised as they are laid, one at a time.

Insects in different groups, including the Odonata (dragonflies and damselflies) and the Hymenoptera (ants, bees, and wasps) practise delayed fertilisation. Among the Odonata, females may mate with multiple males, and store sperm until the eggs are laid. The male may hover above the female during egg-laying (oviposition) to prevent her from mating with other males and replacing his sperm; in some groups such as the darters, the male continues to grasp the female with his claspers during egg-laying, the pair flying around in tandem. Among social Hymenoptera, honeybee queens mate only on mating flights, in a short period lasting some days; a queen may mate with eight or more drones. She then stores the sperm for the rest of her life, perhaps for five years or more.

Fertilisation in fungi

In many fungi (except chytrids), as in some protists, fertilisation is a two step process. First, the cytoplasms of the two gamete cells fuse (called plasmogamy), producing a dikaryotic or heterokaryotic cell with multiple nuclei. This cell may then divide to produce dikaryotic or heterokaryotic hyphae. The second step of fertilisation is karyogamy, the fusion of the nuclei to form a diploid zygote.

In chytrid fungi, fertilisation occurs in a single step with the fusion of gametes, as in animals and plants.

Fertilisation in protists

Fertilisation in protozoa

There are three types of fertilisation processes in protozoa:

  • gametogamy;
  • autogamy;
  • gamontogamy.

Fertilisation in algae

Fertilisation in algae occurs by binary fission. The pseudopodia is first withdrawn and the nucleus starts dividing. When the cytoplasm is divided, the cytoplasm is also divided into two equal parts for each daughter cell. Two daughter cells are produced by one parent cell. It involves the process of mitosis.

Algae, like some land plants, undergo alternation of generations. Some algae are isomorphic, where both the sporophyte (2n) and gameteophyte (n) are the same morphologically. When algae reproduction is described as oogamous, the male and female gametes are different morphologically, where there is a large non-motile egg for female gametes, and the male gamete are uniflagellate (motile). Via the process of syngamy, these will form a new zygote, regenerating the sporophyte generation again.

Fertilisation in fungi-like protists

Fertilisation in fungi. In many fungi (except chytrids), as in some protists, fertilisation is a two step process. ... In chytrid fungi, fertilisation occurs in a single step with the fusion of gametes, as in animals and plants.

Fertilisation and genetic recombination

Meiosis results in a random segregation of the genes that each parent contributes. Each parent organism is usually identical save for a fraction of their genes; each gamete is therefore genetically unique. At fertilisation, parental chromosomes combine. In humans, (2²²)² = 17.6x1012 chromosomally different zygotes are possible for the non-sex chromosomes, even assuming no chromosomal crossover. If crossover occurs once, then on average (4²²)² = 309x1024 genetically different zygotes are possible for every couple, not considering that crossover events can take place at most points along each chromosome. The X and Y chromosomes undergo no crossover events and are therefore excluded from the calculation. The mitochondrial DNA is only inherited from the maternal parent.

The sperm aster and zygote centrosomes

Shortly after the sperm fuse with the egg, the two sperm centrioles form the embryo first centrosome and microtubule aster. The sperm centriole, found near the male pronucleus, recruit egg Pericentriolar material proteins forming the zygote first centrosome. This centrosome nucleates microtubules in the shape of stars called astral microtubules. The microtubules span the whole valium of the egg, allowing the egg pronucleus to use the cables to get to the male pronucleus. As the male and female pronuclei approach each other, the single centrosome split into two centrosomes located in the interphase between the pronuclei. Then the centrosome via the astral microtubules polarizes the genome inside the pronuclei.

Parthenogenesis

Organisms that normally reproduce sexually can also reproduce via parthenogenesis, wherein an unfertilised female gamete produces viable offspring. These offspring may be clones of the mother, or in some cases genetically differ from her but inherit only part of her DNA. Parthenogenesis occurs in many plants and animals and may be induced in others through a chemical or electrical stimulus to the egg cell. In 2004, Japanese researchers led by Tomohiro Kono succeeded after 457 attempts to merge the ova of two mice by blocking certain proteins that would normally prevent the possibility; the resulting embryo normally developed into a mouse.

Allogamy and autogamy

Allogamy, which is also known as cross-fertilisation, refers to the fertilisation of an egg cell from one individual with the male gamete of another.

Autogamy which is also known as self-fertilisation, occurs in such hermaphroditic organisms as plants and flatworms; therein, two gametes from one individual fuse.

Other variants of bisexual reproduction

Some relatively unusual forms of reproduction are:

Gynogenesis: A sperm stimulates the egg to develop without fertilisation or syngamy. The sperm may enter the egg.

Hybridogenesis: One genome is eliminated to produce haploid eggs.

Canina meiosis: (sometimes called "permanent odd polyploidy") one genome is transmitted in the Mendelian fashion, others are transmitted clonally.

Benefits of cross-fertilisation

The major benefit of cross-fertilisation is generally thought to be the avoidance of inbreeding depression. Charles Darwin, in his 1876 book The Effects of Cross and Self Fertilisation in the Vegetable Kingdom (pages 466-467) summed up his findings in the following way.

“It has been shown in the present volume that the offspring from the union of two distinct individuals, especially if their progenitors have been subjected to very different conditions, have an immense advantage in height, weight, constitutional vigour and fertility over the self-fertilised offspring from one of the same parents. And this fact is amply sufficient to account for the development of the sexual elements, that is, for the genesis of the two sexes.”

In addition, it is thought by some, that a long-term advantage of out-crossing in nature is increased genetic variability that promotes adaptation or avoidance of extinction (see Genetic variability).

Algorithmic information theory

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