Haemolymph
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Hemolymph, or haemolymph, is a fluid, analogous to the
blood Blood is a body fluid in the circulatory system of humans and other vertebrates that delivers necessary substances such as nutrients and oxygen to the cells, and transports metabolic waste products away from those same cells. Blood in the c ...
in
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, ...
s, that circulates in the interior of the arthropod (invertebrate) body, remaining in direct contact with the animal's tissues. It is composed of a fluid plasma in which hemolymph cells called
hemocytes A blood cell, also called a hematopoietic cell, hemocyte, or hematocyte, is a cell produced through hematopoiesis and found mainly in the blood. Major types of blood cells include red blood cells (erythrocytes), white blood cells (leukocytes), a ...
are suspended. In addition to hemocytes, the plasma also contains many chemicals. It is the major tissue type of the
open circulatory system The blood circulatory system is a 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 vascular system, tha ...
characteristic of
arthropods 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 ...
(e.g.
arachnid Arachnida () is a class of joint-legged invertebrate animals (arthropods), in the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, harvestmen, camel spiders, whip spiders and vinegaroon ...
s,
crustacean 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 group ...
s and
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
s). In addition, some non-arthropods such as
molluscs 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 taxon, extant species of molluscs are recognized. The number of fossil sp ...
possess a hemolymphatic circulatory system. Oxygen-transport systems were long thought unnecessary in insects, but ancestral and functional
hemocyanin Hemocyanins (also spelled haemocyanins and abbreviated Hc) are proteins that transport oxygen throughout the bodies of some invertebrate animals. These metalloproteins contain two copper atoms that reversibly bind a single oxygen molecule (O2) ...
has been found in the hemolymph. Insect "blood" generally does not carry
hemoglobin Hemoglobin (haemoglobin BrE) (from the Greek word αἷμα, ''haîma'' 'blood' + Latin ''globus'' 'ball, sphere' + ''-in'') (), abbreviated Hb or Hgb, is the iron-containing oxygen-transport metalloprotein present in red blood cells (erythrocyte ...
, although hemoglobin may be present in the tracheal system instead and play some role in respiration.


Method of transport

In the
grasshopper Grasshoppers are a group of insects belonging to the suborder Caelifera. They are among what is possibly the most ancient living group of chewing herbivorous insects, dating back to the early Triassic around 250 million years ago. Grasshopp ...
, the closed portion of the system consists of tubular hearts and an aorta running along the dorsal side of the insect. The hearts pump hemolymph into the sinuses of the
hemocoel The blood circulatory system is a 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 vascular system, tha ...
where exchanges of materials take place. The volume of hemolymph needed for such a system is kept to a minimum by a reduction in the size of the body cavity. The hemocoel is divided into chambers called sinuses. Coordinated movements of the body muscles gradually bring the hemolymph back to the dorsal sinus surrounding the hearts. Between contractions, tiny valves in the wall of the hearts open and allow hemolymph to enter. Hemolymph fills all of the interior (the
hemocoel The blood circulatory system is a 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 vascular system, tha ...
) of the animal's body and surrounds all cells. It contains
hemocyanin Hemocyanins (also spelled haemocyanins and abbreviated Hc) are proteins that transport oxygen throughout the bodies of some invertebrate animals. These metalloproteins contain two copper atoms that reversibly bind a single oxygen molecule (O2) ...
, a
copper Copper is a chemical element with the symbol Cu (from la, cuprum) and atomic number 29. It is a soft, malleable, and ductile metal with very high thermal and electrical conductivity. A freshly exposed surface of pure copper has a pinkis ...
-based protein that turns blue when oxygenated, instead of the
iron Iron () is a chemical element with symbol Fe (from la, ferrum) and atomic number 26. It is a metal that belongs to the first transition series and group 8 of the periodic table. It is, by mass, the most common element on Earth, right in f ...
-based
hemoglobin Hemoglobin (haemoglobin BrE) (from the Greek word αἷμα, ''haîma'' 'blood' + Latin ''globus'' 'ball, sphere' + ''-in'') (), abbreviated Hb or Hgb, is the iron-containing oxygen-transport metalloprotein present in red blood cells (erythrocyte ...
in
red blood cells Red blood cells (RBCs), also referred to as red cells, red blood corpuscles (in humans or other animals not having nucleus in red blood cells), haematids, erythroid cells or erythrocytes (from Greek language, Greek ''erythros'' for "red" and ''k ...
found in vertebrates, giving hemolymph a blue-green color rather than the red color of vertebrate blood. When not oxygenated, hemolymph quickly loses its color and appears grey. The hemolymph of lower arthropods, including most
insects Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (head, thorax and abdomen), three pairs of j ...
, is not used for oxygen transport because these animals respirate through other means, such as tracheas, but it does contain nutrients such as proteins and sugars. Muscular movements by the animal during
locomotion Locomotion means the act or ability of something to transport or move itself from place to place. Locomotion may refer to: Motion * Motion (physics) * Robot locomotion, of man-made devices By environment * Aquatic locomotion * Flight * Locomo ...
can facilitate hemolymph movement, but diverting flow from one area to another is limited. When the
heart The heart is a muscular organ in most animals. This organ pumps blood through the blood vessels of the circulatory system. The pumped blood carries oxygen and nutrients to the body, while carrying metabolic waste such as carbon dioxide t ...
relaxes, hemolymph is drawn back toward the heart through open-ended pores called ostia. Note that the term " ostia" is not specific to insect circulation; it literally means "doors" or "openings", and must be understood in context.


Constituents

Hemolymph can contain nucleating agents that confer extra cellular freezing protection. Such nucleating agents have been found in the hemolymph of insects of several orders, i.e.,
Coleoptera Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describ ...
(beetles),
Diptera Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced ...
(flies), and
Hymenoptera Hymenoptera is a large order (biology), order of insects, comprising the sawfly, sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described, in addition to over 2,000 extinct ones. Many of the species are Par ...
.


Inorganic

Hemolymph is composed of
water Water (chemical formula ) is an inorganic, transparent, tasteless, odorless, and nearly colorless chemical substance, which is the main constituent of Earth's hydrosphere and the fluids of all known living organisms (in which it acts as a ...
,
inorganic In chemistry, an inorganic compound is typically a chemical compound that lacks carbon–hydrogen bonds, that is, a compound that is not an organic compound. The study of inorganic compounds is a subfield of chemistry known as '' inorganic chemist ...
salt Salt is a mineral composed primarily of sodium chloride (NaCl), a chemical compound belonging to the larger class of salts; salt in the form of a natural crystalline mineral is known as rock salt or halite. Salt is present in vast quantitie ...
s (mostly
sodium Sodium is a chemical element with the symbol Na (from Latin ''natrium'') and atomic number 11. It is a soft, silvery-white, highly reactive metal. Sodium is an alkali metal, being in group 1 of the periodic table. Its only stable iso ...
,
chlorine Chlorine is a chemical element with the Symbol (chemistry), symbol Cl and atomic number 17. The second-lightest of the halogens, it appears between fluorine and bromine in the periodic table and its properties are mostly intermediate betwee ...
,
potassium Potassium is the chemical element with the symbol K (from Neo-Latin ''kalium'') and atomic number19. Potassium is a silvery-white metal that is soft enough to be cut with a knife with little force. Potassium metal reacts rapidly with atmosphe ...
,
magnesium Magnesium is a chemical element with the symbol Mg and atomic number 12. It is a shiny gray metal having a low density, low melting point and high chemical reactivity. Like the other alkaline earth metals (group 2 of the periodic ta ...
, and
calcium Calcium is a chemical element with the symbol Ca and atomic number 20. As an alkaline earth metal, calcium is a reactive metal that forms a dark oxide-nitride layer when exposed to air. Its physical and chemical properties are most similar to ...
), and
organic compounds In chemistry, organic compounds are generally any chemical compounds that contain carbon-hydrogen or carbon-carbon bonds. Due to carbon's ability to catenate (form chains with other carbon atoms), millions of organic compounds are known. The s ...
(mostly
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,
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
lipid Lipids are a broad group of naturally-occurring molecules which includes fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids include ...
s). The primary oxygen transporter molecule is
hemocyanin Hemocyanins (also spelled haemocyanins and abbreviated Hc) are proteins that transport oxygen throughout the bodies of some invertebrate animals. These metalloproteins contain two copper atoms that reversibly bind a single oxygen molecule (O2) ...
.


Amino acids

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 ...
hemolymph contains high levels of free amino acids. Most amino acids are present but their relative concentrations vary from species to species. Concentrations of amino acids also vary according to the arthropod stage of development. An example of this is the silkworm and its need for glycine in the production of silk.


Proteins

Proteins present in the hemolymph vary in quantity during the course of development. These proteins are classified by their functions: chroma proteins, protease inhibitors, storage, lipid transport, enzymes, the vitellogenins, and those involved in the immune responses of arthropods. Some hemolymphic proteins incorporate carbohydrates and lipids into the structure.


Other organic constituents

Nitrogen Nitrogen is the chemical element with the symbol N and atomic number 7. Nitrogen is a nonmetal and the lightest member of group 15 of the periodic table, often called the pnictogens. It is a common element in the universe, estimated at se ...
metabolism end products are present in the hemolymph in low concentrations. These include
ammonia Ammonia is an inorganic compound of nitrogen and hydrogen with the formula . A stable binary hydride, and the simplest pnictogen hydride, ammonia is a colourless gas with a distinct pungent smell. Biologically, it is a common nitrogenous was ...
,
allantoin Allantoin is a chemical compound with formula C4H6N4O3. It is also called 5-ureidohydantoin or glyoxyldiureide. It is a diureide of glyoxylic acid. Allantoin is a major metabolic intermediate in most organisms including animals, plants and bacter ...
,
uric acid Uric acid is a heterocyclic compound of carbon, nitrogen, oxygen, and hydrogen with the formula C5H4N4O3. It forms ions and salts known as urates and acid urates, such as ammonium acid urate. Uric acid is a product of the metabolic breakdown of ...
, and
urea Urea, also known as carbamide, is an organic compound with chemical formula . This amide has two amino groups (–) joined by a carbonyl functional group (–C(=O)–). It is thus the simplest amide of carbamic acid. Urea serves an important r ...
. Arthropod
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 are present, most notably the juvenile hormone.
Trehalose Trehalose (from Turkish '' tıgala'' – a sugar derived from insect cocoons + -ose) is a sugar consisting of two molecules of glucose. It is also known as mycose or tremalose. Some bacteria, fungi, plants and invertebrate animals synthesize it ...
can be present and sometimes in great amounts along with
glucose Glucose is a simple sugar with the molecular formula . Glucose is overall the most abundant monosaccharide, a subcategory of carbohydrates. Glucose is mainly made by plants and most algae during photosynthesis from water and carbon dioxide, using ...
. These sugar levels are maintained by the control of hormones. Other
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 can be present. These include
inositol Inositol, or more precisely ''myo''-inositol, is a carbocyclic sugar that is abundant in the brain and other mammalian tissues; it mediates cell signal transduction in response to a variety of hormones, neurotransmitters, and growth factors and ...
,
sugar alcohol Sugar alcohols (also called polyhydric alcohols, polyalcohols, alditols or glycitols) are organic compounds, typically derived from sugars, containing one hydroxyl group (–OH) attached to each carbon atom. They are white, water-soluble solids ...
, hexosamines,
mannitol Mannitol is a type of sugar alcohol used as a sweetener and medication. It is used as a low calorie sweetener as it is poorly absorbed by the intestines. As a medication, it is used to decrease pressure in the eyes, as in glaucoma, and to lower ...
,
glycerol Glycerol (), also called glycerine in British English and glycerin in American English, is a simple triol compound. It is a colorless, odorless, viscous liquid that is sweet-tasting and non-toxic. The glycerol backbone is found in lipids known ...
and those components that are precursors to
chitin Chitin ( C8 H13 O5 N)n ( ) is a long-chain polymer of ''N''-acetylglucosamine, an amide derivative of glucose. Chitin is probably the second most abundant polysaccharide in nature (behind only cellulose); an estimated 1 billion tons of chit ...
. Free
lipid Lipids are a broad group of naturally-occurring molecules which includes fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids include ...
s are present and are used as fuel for flight.


Hemocytes

There are free-floating cells, the hemocytes, within the hemolymph. They play a role in the arthropod
immune system The immune system is a network of biological processes that protects an organism from diseases. It detects and responds to a wide variety of pathogens, from viruses to parasitic worms, as well as cancer cells and objects such as wood splinte ...
. The immune system resides in the hemolymph.


Comparisons to vertebrates

This open system might appear to be inefficient compared to the closed circulatory systems of 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, ...
s, but the two systems have very different demands placed on them. In vertebrates, the circulatory system is responsible for transporting oxygen to all the tissues and removing carbon dioxide from them. It is this requirement that establishes the level of performance demanded of the system. The efficiency of the vertebrate system is far greater than is needed for transporting nutrients, hormones, and so on, whereas in insects, exchange of oxygen and carbon dioxide occurs in the tracheal system. Hemolymph plays no part in the process in most insects. Only in a few insects living in low-oxygen environments are there hemoglobin-like molecules that bind oxygen and transport it to the tissues. Therefore, the demands placed upon the system are much lower. Some arthropods and most molluscs possess the
copper Copper is a chemical element with the symbol Cu (from la, cuprum) and atomic number 29. It is a soft, malleable, and ductile metal with very high thermal and electrical conductivity. A freshly exposed surface of pure copper has a pinkis ...
-containing
hemocyanin Hemocyanins (also spelled haemocyanins and abbreviated Hc) are proteins that transport oxygen throughout the bodies of some invertebrate animals. These metalloproteins contain two copper atoms that reversibly bind a single oxygen molecule (O2) ...
, however, for oxygen transport.


Specialist uses

In some species, hemolymph has other uses than just being a blood analogue. As the insect or arachnid grows, the hemolymph works something like a hydraulic system, enabling the insect or arachnid to expand segments before they are sclerotized. It can also be used hydraulically as a means of assisting movement, such as in arachnid locomotion. Some species of insect or arachnid are able to
autohaemorrhage Autohaemorrhaging, or reflex bleeding, is the action of animals deliberately ejecting blood from their bodies. Autohaemorrhaging has been observed as occurring in two variations. In the first form, blood is squirted toward a predator. The blood of ...
when they are attacked by predators. Queens of the ant genus '' Leptanilla'' are fed with hemolymph produced by the
larvae A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
.Genus ''Leptanilla''
Australian Ants Online
On the other hand, '' Pemphigus spyrothecae'' utilize hemolymph as an adhesive, allowing the species to stick to predators and subsequently attack the predator; it was found that with larger predators, more aphids were stuck after the predator was defeated.


See also

*
Insect physiology Insect physiology includes the physiology and biochemistry of insect organ systems. Although diverse, insects are quite similar in overall design, internally and externally. The insect is made up of three main body regions (tagmata), the head, thor ...
*
Respiratory system of insects An insect's respiratory system is the system with which it introduces respiratory gases to its interior and performs gas exchange. Air enters the respiratory systems of insects through a series of external openings called spiracles. These extern ...


References


Sources

* {{cite book , last=Chapman , first=R.F. , title=The Insects: Structure and Function , url=https://archive.org/details/insectsstructure0000chap , url-access=registration , publisher=Cambridge University Press , edition=4th , year=1998 , location=Cambridge , isbn=978-0-521-57890-5


External links

* ''
Boston Globe ''The Boston Globe'' is an American daily newspaper founded and based in Boston, Massachusetts. The newspaper has won a total of 27 Pulitzer Prizes, and has a total circulation of close to 300,000 print and digital subscribers. ''The Boston Glob ...
'
article
on hemolymph
Blue Squid Blood
Blood Arthropod anatomy Insect anatomy Crustacean anatomy Arachnid anatomy Body fluids