Copepods (; meaning 'oar-feet') are a group of small found in nearly every freshwater and saltwater habitat. Some species are (living in the water column), some are benthos (living on the sediments), several species have Parasitism, and some continental species may live in limnoterrestrial habitats and other wet terrestrial places, such as swamps, under leaf fall in wet forests, bogs, springs, ephemeral ponds, puddles, damp moss, or water-filled recesses of plants (phytotelmata) such as and . Many live underground in marine and freshwater caves, , or stream beds. Copepods are sometimes used as biodiversity indicators.
As with other crustaceans, copepods have a form. For copepods, the egg hatches into a nauplius form, with a head and a tail but no true thorax or abdomen. The larva molts several times until it resembles the adult and then, after more molts, achieves adult development. The nauplius form is so different from the adult form that it was once thought to be a separate species. The metamorphosis had, until 1832, led to copepods being misidentified as or (albeit aquatic ones), or, for parasitic copepods, 'fish lice'.
Free-living copepods of the orders Calanoida, Cyclopoida, and Harpacticoida typically have a short, cylindrical body, with a rounded or beaked head, although considerable variation exists in this pattern. The head is fused with the first one or two thorax segments, while the remainder of the thorax has three to five segments, each with limbs. The first pair of thoracic appendages is modified to form , which assist in feeding. The abdomen is typically narrower than the thorax, and contains five segments without any appendages, except for some tail-like "rami" at the tip. Parasitic copepods (the other seven orders) vary widely in morphology and no generalizations are possible.
Because of their small size, copepods have no need of any heart or circulatory system (the members of the order Calanoida have a heart, but no ), and most also lack . Instead, they absorb oxygen directly into their bodies. Their excretory system consists of maxillary glands.
Some copepods have extremely fast when a predator is sensed, and can jump with high speed over a few millimetres. Many species have surrounded by myelin (for increased conduction speed), which is very rare among (other examples are some and crustaceans like Palaemonidae shrimp and Penaeidae). Even rarer, the myelin is highly organized, resembling the well-organized wrapping found in vertebrates (Gnathostomata). Despite their fast escape response, copepods are successfully hunted by slow-swimming , which approach their prey so gradually, it senses no turbulence, then suck the copepod into their snout too suddenly for the copepod to escape.
Several species are Bioluminescence and able to produce light. It is assumed this is an antipredatory defense mechanism.
Finding a mate in the three-dimensional space of open water is challenging. Some copepod females solve the problem by emitting , which leave a trail in the water that the male can follow. Copepods experience a low Reynolds number and therefore a high relative viscosity. One foraging strategy involves chemical detection of sinking marine snow aggregates and taking advantage of nearby low-pressure gradients to swim quickly towards food sources.
Eggs hatch into nauplius larvae, which consist of a head with a small telson, but no thorax or true abdomen. The nauplius moults five or six times, before emerging as a "copepodid larva". This stage resembles the adult, but has a simple, unsegmented abdomen and only three pairs of thoracic limbs. After a further five moults, the copepod takes on the adult form. The entire process from hatching to adulthood can take a week to a year, depending on the species and environmental conditions such as temperature and nutrition (e.g., egg-to-adult time in the calanoid Parvocalanus crassirostris is ~7 days at but 19 days at .
Because of their smaller size and relatively faster growth rates, and because they are more evenly distributed throughout more of the world's oceans, copepods almost certainly contribute far more to the secondary productivity of the world's oceans, and to the global ocean carbon sink than krill, and perhaps more than all other groups of organisms together. The surface layers of the oceans are believed to be the world's largest carbon sink, absorbing about 2 billion tons of carbon a year, the equivalent to perhaps a third of greenhouse gas, thus reducing their impact. Many planktonic copepods feed near the surface at night, then sink (by changing oils into more density fats) into deeper water during the day to avoid visual predators. Their moulted , feces pellets, and respiration at depth all bring carbon to the deep sea.
About half of the estimated 14,000 described species of copepods are parasite See photograph at Photograph taken by Kerryn Parkinson and Robin McPhee in June 2003. and many have adapted extremely modified bodies for their parasitic lifestyles. They attach themselves to bony fish, sharks, marine mammals, and many kinds of invertebrates such as corals, other crustaceans, molluscs, sponges, and tunicates. They also live as ectoparasites on some freshwater fish.
During the naupliar stage, the copepod host ingests the unicellular dinospore of the parasite. The dinospore is not digested and continues to grow inside the intestinal lumen of the copepod. Eventually, the parasite divides into a multicellular arrangement called a trophont. This trophont is considered parasitic, contains thousands of cells, and can be several hundred micrometers in length. The trophont is greenish to brownish in color as a result of well-defined . At maturity, the trophont ruptures and Blastodinium spp. are released from the copepod anus as free dinospore cells. Not much is known about the dinospore stage of Blastodinium and its ability to persist outside of the copepod host in relatively high abundances.
The copepod Calanus finmarchicus, which dominates the northeastern Atlantic Ocean, has been shown to be greatly infected by this parasite. A 2014 study in this region found up to 58% of collected C. finmarchicus females to be infected. In this study, Blastodinium-infected females had no measurable feeding rate over a 24-hour period. This is compared to uninfected females which, on average, ate 2.93 × 104 cells per day. Blastodinium-infected females of C. finmarchicus exhibited characteristic signs of starvation, including decreased respiration, fecundity, and fecal pellet production. Though photosynthetic, Blastodinium spp. procure most of their energy from organic material in the copepod gut, thus contributing to host starvation. Underdeveloped or disintegrated ovary and decreased fecal pellet size are a direct result of starvation in female copepods. Parasitic infection by Blastodinium spp. could have serious ramifications on the success of copepod species and the function of entire marine ecosystems. Blastodinium parasitism is not lethal, but has negative impacts on copepod physiology, which in turn may alter marine biogeochemical cycles.
Freshwater copepods of the Cyclops genus are the intermediate host of the Guinea worm ( Dracunculus medinensis), the nematode that causes dracunculiasis disease in humans. This disease may be close to being eradicated through efforts by the U.S. Centers for Disease Control and Prevention, the World Health Organization, and the Carter Center.
Copepods are known hosts of Vibrio bacteria, including pathogenic species. The Vibrio attach to the copepod's chitinous carapace, wearing it away to create a niche to stay. They are more protected from ecological stressors when attached to copepods and have an easy dispersal method. Vibrio are not known to infect copepods, but the degradation of the carapace is presumably detrimental to the copepod.
Copepods are infected by a variety of marine fungi including Metschnikowia species, and this can be lethal. They are also parasitized by Cestoda, Isopoda, and many kinds of protist, including Ellobiopsidae, Ciliate, and Apicomplexa.
Copepods have been used successfully in Vietnam to control disease-bearing mosquitoes such as Aedes aegypti that transmit dengue fever and other human parasitic diseases.
The copepods can be added to water-storage containers where the mosquitoes breed. Copepods, primarily of the genera Mesocyclops and Macrocyclops (such as Macrocyclops albidus), can survive for periods of months in the containers, if the containers are not completely drained by their users. They attack, kill, and eat the younger first- and second-instar larvae of the mosquitoes. This biological control method is complemented by community trash removal and recycling to eliminate other possible mosquito-breeding sites. Because the water in these containers is drawn from uncontaminated sources such as rainfall, the risk of contamination by cholera bacteria is small, and in fact no cases of cholera have been linked to copepods introduced into water-storage containers. Trials using copepods to control container-breeding mosquitoes are underway in several other countries, including Thailand and the southern United States. The method, though, would be very ill-advised in areas where the guinea worm is endemic.
When a group of rabbis in Brooklyn, New York, discovered these copepods in the summer of 2004, they triggered such debate in rabbinic circles that some observant Jews felt compelled to buy and install filters for their water. The water was ruled kosher by posek Yisrael Belsky, chief posek of the Orthodox Union and one of the most scientifically literate poskim of his time.Berger, Joseph (November 7, 2004) "The Water's Fine, But Is It Kosher?" , The New York Times Meanwhile, Rabbi Dovid Feinstein, based on the ruling of Rabbi Yosef Shalom Elyashiv - the two widely considered to be the greatest poskim of their time - ruled it was not kosher until filtered. Several major kashrus organizations (e.g OU Kashrus and Star-K) require tap water to have filters.
|
|