The nephridium (plural ''nephridia'') is an
invertebrate
Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...
organ, found in pairs and performing a function similar to the
vertebrate
Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
kidney
The kidneys are two reddish-brown bean-shaped organs found in vertebrates. They are located on the left and right in the retroperitoneal space, and in adult humans are about in length. They receive blood from the paired renal arteries; blood ...
s (which originated from the chordate nephridia). Nephridia remove
metabolic
Metabolism (, from el, μεταβολή ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run cell ...
wastes from an animal's body. Nephridia come in two basic categories: metanephridia and protonephridia. All nephridia- and kidney- having animals belong to the
clade
A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
Nephrozoa
Nephrozoa is a major clade of bilaterians, divided into the protostomes and the deuterostomes, containing almost all animal phyla and over a million extant species. Its sister clade is the Xenacoelomorpha. The Ambulacraria (conventionally deut ...
.
Metanephridia
![Metanephridium](https://upload.wikimedia.org/wikipedia/commons/e/ed/Metanephridium.svg)
A metanephridium (''meta'' = "after") is a type of
excretory
Excretion is a process in which metabolic waste
is eliminated from an organism. In vertebrates this is primarily carried out by the lungs, kidneys, and skin. This is in contrast with secretion, where the substance may have specific tasks aft ...
gland
In animals, a gland is a group of cells in an animal's body that synthesizes substances (such as hormones) for release into the bloodstream (endocrine gland) or into cavities inside the body or its outer surface (exocrine gland).
Structure
De ...
found in many types of
invertebrate
Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...
s such as
annelid
The annelids (Annelida , from Latin ', "little ring"), also known as the segmented worms, are a large phylum, with over 22,000 extant species including ragworms, earthworms, and leeches. The species exist in and have adapted to various ecol ...
s,
arthropod
Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
s and
mollusca
Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000 extant species of molluscs are recognized. The number of fossil species is esti ...
. (In mollusca, it is known as the
Bojanus organ
The organs of Bojanus or Bojanus organs are excretory glands that serve the function of kidneys in some of the molluscs. In other words, these are metanephridia that are found in some molluscs, for example in the bivalves. Some other molluscs hav ...
.)
A metanephridium typically consists of a ciliated funnel opening into the body cavity, or
coelom
The coelom (or celom) is the main body cavity in most animals and is positioned inside the body to surround and contain the digestive tract and other organs. In some animals, it is lined with mesothelium. In other animals, such as molluscs, it r ...
connected to a
duct which may be variously glandularized, folded or expanded (vesiculate) and which typically opens to the
organism
In biology, an organism () is any living system that functions as an individual entity. All organisms are composed of cells (cell theory). Organisms are classified by taxonomy into groups such as multicellular animals, plants, and ...
's exterior. These ciliated tubules pump water carrying surplus
ion
An ion () is an atom or molecule with a net electrical charge.
The charge of an electron is considered to be negative by convention and this charge is equal and opposite to the charge of a proton, which is considered to be positive by conve ...
s,
metabolic waste
Metabolic wastes or excrements are substances left over from metabolic processes (such as cellular respiration) which cannot be used by the organism (they are surplus or toxic), and must therefore be excreted. This includes nitrogen compounds, ...
,
toxin
A toxin is a naturally occurring organic poison produced by metabolic activities of living cells or organisms. Toxins occur especially as a protein or conjugated protein. The term toxin was first used by organic chemist Ludwig Brieger (1849– ...
s from
food
Food is any substance consumed by an organism for nutritional support. Food is usually of plant, animal, or fungal origin, and contains essential nutrients, such as carbohydrates, fats, proteins, vitamins, or minerals. The substance is inge ...
, and useless
hormone
A hormone (from the Greek participle , "setting in motion") is a class of signaling molecules in multicellular organisms that are sent to distant organs by complex biological processes to regulate physiology and behavior. Hormones are required ...
s out of the organism by directing them down funnel-shaped bodies called
nephrostome
The nephrostome is the funnel-like component of a metanephridium. It is always oriented towards the coelom. The nephrostome is covered from the inside with cilia, which push the water, metabolic wastes, unnecessary hormones
A hormone (from th ...
s. This waste is passed out of the organism at the
nephridiopore A nephridiopore is part of the nephridium, an excretory organ found in many organisms, such as flatworms and annelids. Polychaetes typically release their gametes into the water column using nephridiopores.
Nephridia are homologous to nephrons or ...
. The primary
urine
Urine is a liquid by-product of metabolism in humans and in many other animals. Urine flows from the kidneys through the ureters to the urinary bladder. Urination results in urine being excretion, excreted from the body through the urethra.
Cel ...
produced by filtration of blood (or a similarly functioning fluid) is modified into secondary urine through
selective reabsorption Selective reabsorption is the process whereby certain molecules (e.g. ions, glucose and amino acids), after being filtered out of the capillaries along with nitrogenous waste products (i.e. urea) and water in the glomerulus, are reabsorbed from the ...
by the
cells lining the metanephridium.
Saccate metanephridia
The saccate metanephridia are excretory glands which function similarly to the metanephridia. They are found in the
arthropod
Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
s:
coxal gland
The coxal gland is a gland found in some arthropods, for collecting and excreting urine. They are found in all arachnids (with the exception of some Acari), and in other chelicerates, such as horseshoe crabs. The coxal gland is thought to be homo ...
s of arachnids, antennal (or green) glands and maxillary glands of crustaceans, etc.
The saccate metanephridia filter the fluid of the
hemocoel
The blood circulatory system is a organ system, system of organs that includes the heart, blood vessels, and blood which is circulated throughout the entire body of a human or other vertebrate. It includes the cardiovascular system, or vascula ...
, as opposed to the metanephridia which filter
coelomic fluid. In a saccate metanephridium, there is a ciliated funnel covered with a membrane that helps to filter the hemocoel of
heavy
Heavy may refer to:
Measures
* Heavy (aeronautics), a term used by pilots and air traffic controllers to refer to aircraft capable of 300,000 lbs or more takeoff weight
* Heavy, a characterization of objects with substantial weight
* Heavy, ...
particles (such as
protein
Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respo ...
s and
carbohydrate
In organic chemistry, a carbohydrate () is a biomolecule consisting of carbon (C), hydrogen (H) and oxygen (O) atoms, usually with a hydrogen–oxygen atom ratio of 2:1 (as in water) and thus with the empirical formula (where ''m'' may or ma ...
s) before the fluid even enters the funnel. Inside the funnel, the fluid is further processed through selective reabsorption, and eventually
excreted
Excretion is a process in which metabolic waste
is eliminated from an organism. In vertebrates this is primarily carried out by the lungs, kidneys, and skin. This is in contrast with secretion, where the substance may have specific tasks after lea ...
from the
nephridiopore A nephridiopore is part of the nephridium, an excretory organ found in many organisms, such as flatworms and annelids. Polychaetes typically release their gametes into the water column using nephridiopores.
Nephridia are homologous to nephrons or ...
.
In
Crustacea
Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean gro ...
, the saccate metanephridia are associated with the antennae and form the
antennal gland
Antennae ( antenna), sometimes referred to as "feelers", are paired appendages used for sensing in arthropods.
Antennae are connected to the first one or two segments of the arthropod head. They vary widely in form but are always made of one o ...
. In freshwater
crustacea
Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean gro ...
, the saccate metanephridia are especially large due to their role in osmoregulation; crustacea must remove large amounts of water from the tissues, as the cells are
hypertonic
In chemical biology, tonicity is a measure of the effective osmotic pressure gradient; the water potential of two solutions separated by a partially-permeable cell membrane. Tonicity depends on the relative concentration of selective membrane- ...
to the surrounding water.
Protonephridia
![Flamecell](https://upload.wikimedia.org/wikipedia/commons/f/f7/Flamecell.jpg)
A protonephridium (''proto'' = "first") is found in the phyla
Platyhelminthes
The flatworms, flat worms, Platyhelminthes, or platyhelminths (from the Greek language, Greek πλατύ, ''platy'', meaning "flat" and ἕλμινς (root: ἑλμινθ-), ''helminth-'', meaning "worm") are a Phylum (biology), phylum of relati ...
,
Nemertea
Nemertea is a phylum of animals also known as ribbon worms or proboscis worms, consisting of 1300 known species. Most ribbon worms are very slim, usually only a few millimeters wide, although a few have relatively short but wide bodies. Many h ...
,
Rotifera
The rotifers (, from the Latin , "wheel", and , "bearing"), commonly called wheel animals or wheel animalcules, make up a phylum (Rotifera ) of microscopic and near-microscopic pseudocoelomate animals.
They were first described by Rev. John Ha ...
and
Chordata
A chordate () is an animal of the phylum Chordata (). All chordates possess, at some point during their larval or adult stages, five synapomorphies, or primary physical characteristics, that distinguish them from all the other taxa. These fiv ...
(
lancelet
The lancelets ( or ), also known as amphioxi (singular: amphioxus ), consist of some 30 to 35 species of "fish-like" benthic filter feeding chordates in the order Amphioxiformes. They are the modern representatives of the subphylum Cephalochorda ...
s). They have the same anatomy as the
metanephridia
The nephridium (plural ''nephridia'') is an invertebrate organ, found in pairs and performing a function similar to the vertebrate kidneys (which originated from the chordate nephridia). Nephridia remove metabolic wastes from an animal's body. Ne ...
but with the internal ciliated funnel blocked by terminal cells: either a ''
flame cell
A flame cell is a specialized excretory cell found in the simplest freshwater invertebrates, including flatworms, rotifers and nemerteans; these are the simplest animals to have a dedicated excretory system. Flame cells function like a kidney, ...
s'' (if ciliated) or a ''
solenocyte
In biology, solenocytes are flagellated cells associated with excretion, osmoregulation and ionoregulation in many animals and in some chordates under the sub-phylum Cephalochordata.
These are the cells which form subtypes of protonephridium alon ...
s'' (if flagellated). Thus their
tubule
In biology, a tubule is a general term referring to small tube or similar type of structure. Specifically, tubule can refer to:
* a small tube or fistular structure
* a minute tube lined with glandular epithelium
* any hollow cylindrical body stru ...
s lack internal openings, while retaining their opening to the organism's exterior. They function in
osmoregulation
Osmoregulation is the active regulation of the osmotic pressure of an organism's body fluids, detected by osmoreceptors, to maintain the homeostasis of the organism's water content; that is, it maintains the fluid balance and the concentration o ...
(ionoregulation).
Each terminal cell has one or more
cilia
The cilium, plural cilia (), is a membrane-bound organelle found on most types of eukaryotic cell, and certain microorganisms known as ciliates. Cilia are absent in bacteria and archaea. The cilium has the shape of a slender threadlike projecti ...
and their beating inside the protonephridial tube creates an outward going current and hence a partial
pressurization
{{Wiktionary
Pressurization or pressurisation is the application of pressure in a given situation or environment.
Industrial
Industrial equipment is often maintained at pressures above or below atmospheric.
Atmospheric
This is the process by ...
in the blind of the tube. Because of this, pressurization drives
waste fluids from the inside of the animal, and they are pulled through small perforations in the terminal cells and into the protonephridium. The perforations in the terminal cell are large enough for small
molecule
A molecule is a group of two or more atoms held together by attractive forces known as chemical bonds; depending on context, the term may or may not include ions which satisfy this criterion. In quantum physics, organic chemistry, and bioch ...
s to pass, but larger
protein
Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respo ...
s are retained within the animal. From the bottom of the protonephridium the solutes are led through the tube, formed by the canal cells, and exits the animal from a small opening formed by the
nephridiopore A nephridiopore is part of the nephridium, an excretory organ found in many organisms, such as flatworms and annelids. Polychaetes typically release their gametes into the water column using nephridiopores.
Nephridia are homologous to nephrons or ...
.
Selective reabsorption of useful molecules by the canal cells occurs as the solutes pass down the tubule. Protonephridia are generally found in basal organisms such as
flatworm
The flatworms, flat worms, Platyhelminthes, or platyhelminths (from the Greek πλατύ, ''platy'', meaning "flat" and ἕλμινς (root: ἑλμινθ-), ''helminth-'', meaning "worm") are a phylum of relatively simple bilaterian, unsegment ...
s. Protonephridia likely first arose as a way to cope with a hypotonic environment by removing excess water from the organism (
osmoregulation
Osmoregulation is the active regulation of the osmotic pressure of an organism's body fluids, detected by osmoreceptors, to maintain the homeostasis of the organism's water content; that is, it maintains the fluid balance and the concentration o ...
). Their use as excretory and ionoregulatory structures likely arose secondarily.
These are excretory systems in phyla
Platyhelminthes
The flatworms, flat worms, Platyhelminthes, or platyhelminths (from the Greek language, Greek πλατύ, ''platy'', meaning "flat" and ἕλμινς (root: ἑλμινθ-), ''helminth-'', meaning "worm") are a Phylum (biology), phylum of relati ...
and are also called blind tubules. These tubules bear a tuft of
cilia
The cilium, plural cilia (), is a membrane-bound organelle found on most types of eukaryotic cell, and certain microorganisms known as ciliates. Cilia are absent in bacteria and archaea. The cilium has the shape of a slender threadlike projecti ...
or
flagellum
A flagellum (; ) is a hairlike appendage that protrudes from certain plant and animal sperm cells, and from a wide range of microorganisms to provide motility. Many protists with flagella are termed as flagellates.
A microorganism may have f ...
. An organ of excretion in flatworms: a hollow cup-shaped cell containing a bunch of cilia or flagellum, whose movement draws in waste products and wafts them to the outside through a connecting tubule.
External links
*http://www.biology.ualberta.ca/courses.hp/zool250/animations/Excretion.swf
* Baeumler N., Haszprunar G. & Ruthensteiner B. (2012). "Development of the excretory system in a polyplacophoran mollusc: stages in metanephridial system development". ''
Frontiers in Zoology
''Frontiers in Zoology'' is a peer-reviewed open access scientific journal covering all aspects of zoology. It was established in 2004 and is published by BioMed Central on behalf of the . The editors-in-chief are
Jürgen Heinze (University of Re ...
'' 9: 23. .
{{Gastropod anatomy
Annelid anatomy
Arthropod anatomy
Mollusc anatomy
Platyhelminth anatomy