Kidney development, or nephrogenesis, describes the embryologic origins of the kidney, a major organ in the urinary system. This article covers a 3 part developmental process that is observed in most reptiles, birds and mammals, including humans. Nephrogenesis is often considered in the broader context of the development of the urinary and reproductive organs.
The development of the kidney proceeds through a series of successive phases, each marked by the development of a more advanced kidney: the archinephros, pronephros, mesonephros, and metanephros. [1] The pronephros is the most immature form of kidney, while the metanephros is most developed. The metanephros persists as the definitive adult kidney.[ citation needed ]
The archinephros is considered as hypothetical or primitive kidney.
The pronephros develops in the cervical region of the embryo. During approximately day 22 of human gestation, the paired pronephri appears towards the cranial end of the intermediate mesoderm. In this region, epithelial cells arrange themselves in a series of tubules called nephrotomes and join laterally with the pronephric duct. This duct is fully contained within the embryo and thus cannot excrete filtered material outside the embryo; therefore the pronephros is considered nonfunctional in humans.
The development of the pronephric duct proceeds in a cranial-to-caudal direction. As it elongates caudally, the pronephric duct induces nearby intermediate mesoderm in the thoracolumbar area to become epithelial tubules called mesonephric tubules. Each mesonephric tubule receives a blood supply from a branch of the aorta, ending in a capillary tuft analogous to the glomerulus of the definitive nephron. The mesonephric tubule forms a capsule around the capillary tuft, allowing for filtration of blood. This filtrate flows through the mesonephric tubule and is drained into the continuation of the pronephric duct, now called the mesonephric duct or Wolffian duct. The nephrotomes of the pronephros degenerate while the mesonephric duct extends towards the most caudal end of the embryo, ultimately attaching to the cloaca. The mammalian mesonephros is similar to the kidneys of aquatic amphibians and fishes.
During the fifth week of gestation, the mesonephric duct develops an outpouching, the ureteric bud, near its attachment to the cloaca. This bud, also called the metanephrogenic diverticulum, grows posteriorly and towards the head of the embryo. The elongated stalk of the ureteric bud, called the metanephric duct, later forms the ureter. As the cranial end of the bud extends into the intermediate mesoderm, it undergoes a series of branchings to form the collecting duct system of the kidney. It also forms the major and minor calyces and the renal pelvis.
The portion of undifferentiated intermediate mesoderm in contact with the tips of the branching ureteric bud is known as the metanephrogenic blastema. Signals released from the ureteric bud induce the differentiation of the metanephrogenic blastema into the renal tubules. As the renal tubules grow, they come into contact and join with connecting tubules of the collecting duct system, forming a continuous passage for flow from the renal tubule to the collecting duct. Simultaneously, precursors of vascular endothelial cells begin to take their position at the tips of the renal tubules. These cells differentiate into the cells of the definitive glomerulus.
In humans, all of the branches of the ureteric bud and the nephronic units have been formed by 32 to 36 weeks of gestation. However, these structures are not yet mature, and will continue to mature after birth. Once matured, humans have an estimated two million nephrons (approximately 1,000,000 per kidney) but this number is highly variable ranging widely from approximately 200,000 to over 2.5 million per kidney. [2]
After inducing the metanephric mesenchyme the lower portions of the nephric duct will migrate caudally (downward) and connect with the bladder, thereby forming the ureters. The ureters will carry urine from the kidneys to the bladder for excretion from the fetus into the amniotic sac. As the fetus develops, the torso elongates and the kidneys rotate and migrate upwards within the abdomen which causes the length of the ureters to increase.
In humans, the kidneys are two reddish-brown bean-shaped blood-filtering organs that are a multilobar, multipapillary form of mammalian kidneys, usually without signs of external lobulation. They are located on the left and right in the retroperitoneal space, and in adult humans are about 12 centimetres in length. They receive blood from the paired renal arteries; blood exits into the paired renal veins. Each kidney is attached to a ureter, a tube that carries excreted urine to the bladder.
The ureters are tubes made of smooth muscle that propel urine from the kidneys to the urinary bladder. In a human adult, the ureters are usually 20–30 cm (8–12 in) long and around 3–4 mm (0.12–0.16 in) in diameter. The ureter is lined by urothelial cells, a type of transitional epithelium, and has an additional smooth muscle layer that assists with peristalsis in its lowest third.
The nephron is the minute or microscopic structural and functional unit of the kidney. It is composed of a renal corpuscle and a renal tubule. The renal corpuscle consists of a tuft of capillaries called a glomerulus and a cup-shaped structure called Bowman's capsule. The renal tubule extends from the capsule. The capsule and tubule are connected and are composed of epithelial cells with a lumen. A healthy adult has 1 to 1.5 million nephrons in each kidney. Blood is filtered as it passes through three layers: the endothelial cells of the capillary wall, its basement membrane, and between the foot processes of the podocytes of the lining of the capsule. The tubule has adjacent peritubular capillaries that run between the descending and ascending portions of the tubule. As the fluid from the capsule flows down into the tubule, it is processed by the epithelial cells lining the tubule: water is reabsorbed and substances are exchanged ; first with the interstitial fluid outside the tubules, and then into the plasma in the adjacent peritubular capillaries through the endothelial cells lining that capillary. This process regulates the volume of body fluid as well as levels of many body substances. At the end of the tubule, the remaining fluid—urine—exits: it is composed of water, metabolic waste, and toxins.
The mesonephric duct, also known as the Wolffian duct, archinephric duct, Leydig's duct or nephric duct, is a paired organ that develops in the early stages of embryonic development in humans and other mammals. It is an important structure that plays a critical role in the formation of male reproductive organs. The duct is named after Caspar Friedrich Wolff, a German physiologist and embryologist who first described it in 1759.
The collecting duct system of the kidney consists of a series of tubules and ducts that physically connect nephrons to a minor calyx or directly to the renal pelvis. The collecting duct participates in electrolyte and fluid balance through reabsorption and excretion, processes regulated by the hormones aldosterone and vasopressin.
The genitourinary system, or urogenital system, are the organs of the reproductive system and the urinary system. These are grouped together because of their proximity to each other, their common embryological origin and the use of common pathways, like the male urethra. Also, because of their proximity, the systems are sometimes imaged together.
The development of the urinary system begins during prenatal development, and relates to the development of the urogenital system – both the organs of the urinary system and the sex organs of the reproductive system. The development continues as a part of sexual differentiation.
The mesonephros is one of three excretory organs that develop in vertebrates. It serves as the main excretory organ of aquatic vertebrates and as a temporary kidney in reptiles, birds, and mammals. The mesonephros is included in the Wolffian body after Caspar Friedrich Wolff who described it in 1759.
Pronephros is the most basic of the three excretory organs that develop in vertebrates, corresponding to the first stage of kidney development. It is succeeded by the mesonephros, which in fish and amphibians remains as the adult kidney. In amniotes, the mesonephros is the embryonic kidney and a more complex metanephros acts as the adult kidney. Once a more advanced kidney forms, the previous version typically degenerates by apoptosis or becomes part of the male reproductive system.
Intermediate mesoderm or intermediate mesenchyme is a narrow section of the mesoderm located between the paraxial mesoderm and the lateral plate of the developing embryo. The intermediate mesoderm develops into vital parts of the urogenital system.
The ureteric bud, also known as the metanephric diverticulum, is a protrusion from the mesonephric duct during the development of the urinary and reproductive organs. It later develops into a conduit for urine drainage from the kidneys, which, in contrast, originate from the metanephric blastema.
The metanephrogenic blastema or metanephric blastema is one of the two embryological structures that give rise to the kidney, the other being the ureteric bud.
In earlier conceptions of kidney biology, the nephrotome was a section of the mesoderm that gives rise to the pronephros and eventually to the rest of the kidney. Older texts describe the pronephros as forming through the fusion of multiple nephrotomes.
Human embryonic development or human embryogenesis is the development and formation of the human embryo. It is characterised by the processes of cell division and cellular differentiation of the embryo that occurs during the early stages of development. In biological terms, the development of the human body entails growth from a one-celled zygote to an adult human being. Fertilization occurs when the sperm cell successfully enters and fuses with an egg cell (ovum). The genetic material of the sperm and egg then combine to form the single cell zygote and the germinal stage of development commences. Embryonic development in the human, covers the first eight weeks of development; at the beginning of the ninth week the embryo is termed a fetus. The eight weeks has 23 stages.
The development of the reproductive system is the part of embryonic growth that results in the sex organs and contributes to sexual differentiation. Due to its large overlap with development of the urinary system, the two systems are typically described together as the urogenital or genitourinary system.
Protein odd-skipped-related 1 is a transcription factor that in humans is encoded by the OSR1 gene. The OSR1 and OSR2 transcription factors participate in the normal development of body parts such as the kidney.
Duplicated ureter or duplex collecting system is a congenital condition in which the ureteric bud, the embryological origin of the ureter, splits, resulting in two ureters draining a single kidney. It is the most common renal abnormality, occurring in approximately 1% of the population.
The archinephros, or holonephros, is a primitive kidney that has been retained by the larvae of hagfish and some caecilians. A recent author has referred to this structure as "the hypothetical primitive kidney of ancestral vertebrates". In the earliest vertebrates, this structure potentially extended the entire length of the body and consisted of paired segmental structures which drained via a pair of archinephrenic ducts into the cloaca. The entire structure arises from the nephric ridge, which in higher animal embryos gives rise to nephrotomes and the pronephroi at around 4 weeks gestation in humans. The pronephroi are supplanted by mesonephroi and finally by definitive kidneys, the metanephroi, by around 5 weeks gestation. The archinephros is nonfunctional in humans and other mammals.
The kidneys are a pair of organs of the excretory system in vertebrates, which maintains the balance of water and electrolytes in the body (osmoregulation), filters the blood, removes metabolic waste products, and in many vertebrates also produces hormones and maintains blood pressure. In healthy vertebrates, the kidneys maintain homeostasis of extracellular fluid in the body. When the blood is being filtered, the kidneys form urine, which consists of water and excess or unnecessary substances, the urine is then excreted from the body through other organs, which in vertebrates, depending on the species, may include the ureter, urinary bladder, cloaca, and urethra.
The mammalian kidneys are a pair of excretory organs of the urinary system of mammals. They are a type of metanephric kidney. The kidneys in mammals are usually bean-shaped and located behind the peritoneum (retroperitoneally) on the back (dorsal) wall of the body. Each kidney consists of a renal capsule, peripheral cortex, internal medulla, forming papilla with its innermost part, calices, and renal pelvis, although the calices or renal pelvis may be absent in some species. Urine is excreted from the kidney through the ureter. Nitrogen-containing waste products are excreted by the kidneys in mammals mainly in the form of urea.