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The glomerulus (/ɡləˈmɛr(j)ələs, ɡl-/), plural glomeruli, is a network of capillaries known as a tuft, located at the beginning of a nephron in the kidney. The tuft is structurally supported by intraglomerular mesangial cells. The blood is filtered across the capillary walls of this tuft through the glomerular filtration barrier, which yields its filtrate of water and soluble substances to a cup-like sac known as Bowman's capsule. The filtrate then enters the renal tubule, of the nephron.[1]

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Glomerular Physiology.png
Looped capillaries of glomerulus between the arterioles
Details
Precursor Metanephric blastema
Identifiers
Latin glomerulus renalis
MeSH D007678
FMA 15624
Anatomical terminology

The glomerulus receives its blood supply from an afferent arteriole of the renal arterial circulation. Unlike most capillary beds, the glomerular capillaries exit into efferent arterioles rather than venules. The resistance of the efferent arterioles causes sufficient hydrostatic pressure within the glomerulus to provide the force for ultrafiltration.

The glomerulus and its surrounding Bowman's capsule constitute a renal corpuscle, the basic filtration unit of the kidney.[2] The rate at which blood is filtered through all of the glomeruli, and thus the measure of the overall renal function, is the glomerular filtration rate (GFR).

Contents

StructureEdit

The glomerulus is a tuft of small blood vessels called capillaries located within Bowman's capsule within the kidney.[2] Glomerular mesangial cells structurally support the tufts. Blood enters the capillaries of the glomerulus by a single arteriole called an afferent arteriole and leaves by an efferent arteriole.[3] The capillaries consist of a tube lined by endothelial cells with a central lumen. The walls have a unique structure: there are pores between the cells that allow water and soluble substances to exit, and after passing through the glomerular basement membrane, and between podocyte foot processes, enter the capsule as ultrafiltrate.

Blood supplyEdit

 
Figure 2: (a) Diagram of the juxtaglomerular apparatus: it has specialized cells working as a unit which monitor the sodiujuxtaglomerular apparatus: it has three types of specm content of the fluid in the distal convoluted tubule (not labelled - it's the tubule on the left) and adjust the glomerular filtration rate and the rate of renin release. (b) Micrograph showing the glomerulus and surrounding structures.

LiningEdit

 
Scanning electron microscope view of the inner surface of an opened (broken) capillary with fenestrae visible.(100,000x magnification)

Capillaries of the glomerulus are lined by endothelial cells. These contain numerous pores - also called fenestrae - 50–100 nm in diameter.[4] Unlike those of other capillaries with fenestrations, these fenestrations are not spanned by diaphragms.[4] They allow for the filtration of fluid, blood plasma solutes and protein, at the same time preventing the filtration of red blood cells, white blood cells, and platelets.

The glomerulus has a glomerular basement membrane (GBM) consisting mainly of laminins, type IV collagen, agrin and nidogen, which are synthesized and secreted by both endothelial cells and podocytes: thus the GBM is sandwiched between the glomerular capillaries and the podocytes. The GBM is 250–400 nm in thickness, which is thicker than basement membranes of other tissue. It is a barrier to blood proteins such as albumin and globulin.[5]

The part of the podocyte in contact with the GBM is called a podocyte foot process or pedicle (Fig. 3): there are gaps between the foot processes through which the filtrate flows into Bowman's space of the capsule.[4] . The space between adjacent podocyte foot processes is spanned by slit diaphragms consisting of a mat of proteins, including podocin and nephrin. In addition, foot processes have a negatively charged coat (glycocalyx) that repels negatively charged molecules such as serum albumin.

Intraglomerular mesangial cellsEdit

The space between the capillaries of a glomerulus is occupied by intraglomerular mesangial cells. They are not part of the filtration barrier but are specialized pericytes that participate in the regulation of the filtration rate by contracting or expanding: they contain actin and myosin filaments to accomplish this. Some mesangial cels are in physical contact with capillaries, others are in physical contact with podocytes. There is two-way chemical cross talk among the mesangial cells, the capillaries, and the podocytes to fine-tune the GFR.

Intrarenal circulationEdit

 
(Fig. 4) Diagram (with labels) of the circulation related to a single glomerulus, associated tubule, and collecting system.

Blood exits the glomerular capillaries by an efferent arteriole instead of a venule, as is seen in the majority of capillary systems (Fig. 4). [3] This provides tighter control over the blood flow through the glomerulus, since arterioles dilate and constrict more readily than venules, owing to their thick circular smooth muscle layer (tunica media). The blood exiting the efferent arteriole enters a renal venule, which in turn enters a renal interlobular vein and then into the renal vein.

Cortical nephrons near the corticomedullary junction (15% of all nephrons) are called juxtamedullary nephrons. The blood exiting the efferent arterioles of these nephrons enter the vasa recta, which are straight capillary branches that deliver blood to the renal medulla. These vasa recta run adjacent to the descending and ascending loop of Henle, and participate in the maintenance of the medullary countercurrent exchange system.

Filtrate drainageEdit

The filtrate that has passed through the three-layered filtration unit enters Bowman's space. From there, it flows into the renal tubule - the nephron - which follows a U-shaped path to the collecting ducts, finally exiting into a renal calyx as urine.

FunctionEdit

FiltrationEdit

 
Scheme of filtration barrier (blood-urine) in the kidney. A. The endothelial cells of the glomerulus; 1. pore (fenestra).
B. Glomerular basement membrane: 1. lamina rara interna 2. lamina densa 3. lamina rara externa
C. Podocytes: 1. enzymatic and structural proteins 2. filtration slit 3. diaphragma

The main function of the glomerulus is to filter plasma to produce glomerular filtrate, which passes down the length of the nephron tubule to form urine. The rate at which the glomerulus produces filtrate from plasma (the glomerular filtration rate) is much higher than in systemic capillaries because of the particular anatomical characteristics of the glomerulus. Unlike systemic capillaries, which receive blood from high-resistance arterioles and drain to low-resistance venules, glomerular capillaries are connected in both ends to high-resistance arterioles: the afferent arteriole, and the efferent arteriole. This arrangement of two arterioles in series determines the high hydrostatic pressure on glomerular capillaries, which is one of the forces that favour filtration to the Bowman's capsule.[6]

If a substance has passed through the glomerular capillary endothelial cells, glomerular basement membrane, and podocytes, then it enters the lumen of the tubule and is known as glomerular filtrate. Otherwise, it exits the glomerulus through the efferent arteriole and continues circulation as discussed below and as shown on the picture.

PermeabilityEdit

The structures of the layers determine their permeability-selectivity (permselectivity). The factors that influence permselectivity are the negative charge of the basement membrane and the podocytic epithelium, and the effective pore size of the glomerular wall (8 nm). As a result, large and/or negatively charged molecules will pass through far less frequently than small and/or positively charged ones.[7] For instance, small ions such as sodium and potassium pass freely, while larger proteins, such as hemoglobin and albumin have practically no permeability at all.

The oncotic pressure on glomerular capillaries is one of the forces that resist filtration. Because large and negatively charged proteins have a low permeability, they cannot filtrate easily to the Bowman's capsule. Therefore, the concentration of these proteins tends to increase as the glomerular capillaries filtrate plasma, increasing the oncotic pressure along the length of a glomerular capillary.[6]

Starling equationEdit

The rate of filtration from the glomerulus to the Bowman's capsule is determined (as in systemic capillaries) by the Starling equation:[6]

 

Regulation of blood pressureEdit

The walls of the afferent arteriole contain specialized smooth muscle cells that synthesize renin. These juxtaglomerular cells play a major role in the renin–angiotensin system, which helps regulate blood volume and pressure.

Clinical significanceEdit

Damage to the glomerulus by disease can allow passage through the glomerular filtration barrier of red blood cells, white blood cells, platelets, and blood proteins such as albumin and globulin. Underlying causes for glomerular injury can be inflammatory, toxic or metabolic. [8] These can be seen in the urine (urinalysis) on microscopic and chemical (dipstick) examination. Examples are diabetic kidney disease, glomerulonephritis, and IgA nephropathy.

HistoryEdit

In 1666, Italian biologist and anatomist Marcello Malpighi first described the glomeruli and demonstrated their continuity with the renal vasculature (281,282). About 175 years later, surgeon and anatomist William Bowman elucidated in detail the capillary architecture of the glomerulus and the continuity between its surrounding capsule and the proximal tubule.[9]

Additional imagesEdit

ReferencesEdit

  1. ^ Pavenstädt H,; Kriz W,; Kretzler M. "Cell biology of the glomerular podocyte". doi:10.1152/physrev.00020.2002. PMID 12506131. 
  2. ^ a b WHEATER 2006, p. 304.
  3. ^ a b WHEATER 2006, p. 307.
  4. ^ a b c WHEATER 2006, p. 310.
  5. ^ Suh, JH; Miner, JH (2013). "The glomerular basement membrane as a barrier to albumin". Nature Reviews. Nephrology. 9 (8): 470–477. doi:10.1038/nrneph.2013.109. PMC 3839671 . PMID 23774818. 
  6. ^ a b c Boron, WF.; Boulapep, EL. (2012). Medical Physiology (2nd ed.). Philadelphia: Saunders. pp. 771, 774. ISBN 978-1437717532. 
  7. ^ Guyton, Arthur C.; Hall, John E. (2006). Textbook of Medical Physiology. Philadelphia: Elsevier Saunders. pp. 316–317. ISBN 0-7216-0240-1. 
  8. ^ Wiggins, RC. "The spectrum of podocytopathies: a unifying view of glomerular diseases". doi:10.1038/sj.ki.5002222. PMID 17410103. 
  9. ^ "lippicotts histology for pathologesits; satcey e. mills

SourcesEdit

  • Hall, Arthur C. Guyton, John E. (2005). Textbook of medical physiology (11th ed.). Philadelphia: W.B. Saunders. p. Chapter 26. ISBN 978-0-7216-0240-0. 
  • Deakin, Barbara Young ... [] ; drawings by Philip J.; et al. (2006). Wheater's functional histology : a text and colour atlas (5th ed.). [Edinburgh?]: Churchill Livingstone/Elsevier. p. Chapter 16. ISBN 978-0-443068508.