James H. Thorp, ... Lawrence L. Lovell, in Thorp and Covich's Freshwater Invertebrates (Fourth Edition), 2016

Terminology and also Morphology

Knowledge of only 5 morphological terms is forced at this higher level: annuli, chaetae (also spelled chaete), clitellum, parapodia, and peristomium. Annuli are ring-like human body segments. Chaetae room bristles which have the right to have different shapes, relying on the taxon. A clitellum is a modification of the epidermis, which establishes as an conveniently visible, glandular girdle partly behind the female pores; the secretes a cocoon in i m sorry eggs are laid. Parapodia space paired, unjointed lateral appendages uncovered in polychaete worms, i m sorry are regularly fleshy (especially in marine polychaetes) and used because that locomotion, respiration, and other functions. The peristomium is the 2nd body segment that annelids. It may be combined with the anterior segment, or prostomium.

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URL: https://www.jajalger2018.org/science/article/pii/B9780123850287000123

Comparative Reproduction

Susan D. Hill, ... Daniel H. Shain, in Encyclopedia the Reproduction (Second Edition), 2018

Cocoon Production

In coordination with egg laying, clitellates secrete a proteinaceous cocoon that provides a microenvironment for embryonic advancement (Coleman and Shain, 2009). The procedure initiates through the proliferation the clitellum-specific granular cell that differentiate into 2 cell types, among which provides fibrous protein the builds the cocoon wall and another that secretes a glue-like substance to seal the cocoon ends. Number of million granules of each material type (i.e., cocoon wall, glue) are necessary to construct a single cocoon. Secreted wall surface granules initially self-assemble to type a sheath around the clitellum. Upon the relax of eggs and cocoon liquid from the mrs pore into the sheath, the worm pulls its head through while simultaneously sealing both ends of the sheath v granulated structures called opercula (Sawyer, 1986). In some leeches (e.g., Helobdella), the cocoon is secreted native a ring the granular cells neighboring the female pore and also forms a sac the is sealed with a solitary operculum. Frequently only a couple of cocoons are secreted end the course of numerous hours, but up come ∼50 have actually been report in some worms (e.g., Myzobdella lugubris). The variety of eggs deposited into each cocoon ranges from one to much more than 100, depending on the species.

Clitellate cocoons display screen a variety of morphologies (Fig. 5) yet fall into three basic categories: hard-shelled, membranous and gelatinous (Siddall and Burreson, 1996). These are differentiated mostly by the thickness of the fibrous cocoon wall. In part terrestrial taxa (e.g., Hirudo, Haemopis), a spongy procession surrounds the cocoon and appears to protect against dessication by trapping water droplets. The contents of clitellate cocoons display screen extraordinary physiochemical nature (e.g., thermal, chemical resilience), do them valuable resources together biomaterials and also bioadhesives.

Fredric R. Govedich, william E. Moser, in Thorp and also Covich's Freshwater Invertebrates (Fourth Edition), 2015


All leeches are hermaphrodites, mirroring either protandry or cosexuality (Davies and Singhal, 1988), v reciprocal cross-fertilization as the general rule. Fertilization, i m sorry is internal, is achieved in most Glossiphoniidae and also all Piscicolidae and Erpobdellidae by it is registered a spermatophore to the partner’s body. Spermatozoa permeate the body wall and make their means to the ovisacs via the coelomic sinuses (Sawyer, 1986; Govedich et al., 2009). The clitellar an ar is the most constant site because that the deposition of spermatophores, yet they deserve to be attached come any component of the body. In some types of Piscicolidae, a devoted region exists for reception that spermatophores; fertilization is successful just by spermatophores deposit there. In Hirudiniformes, reciprocal inner fertilization is brought about by the insertion of an eversible prick into the partner’s vagina.

Once fertilization occurs, eggs are deposited into a cocoon secreted by the clitellum (Figure 23.10). There have the right to be a considerable hold-up between copulation and cocoon deposition. Because that example, in field populations that H. Stagnalis, copulation occurs in the fall and also cocoon deposition takes ar in the feather (Davies and Reynoldson, 1976). It has actually been widely assumed the the existence of a clearly shows clitellum closely parallels mature of the female reproductive system; and in nonreproductive leeches, the clitellar glands space barely distinguishable from the epithelial cell (Fernandez et al., 1992). The clitellum is particularly prominent in Erpobdellidae. Biernacka and Davies (1995) verified that sexual maturity that E. Obscura can not be judged by the existence or lack of an externally clearly shows clitellum. Although a high relationship of the populace exhibited a clearly shows clitellum at part time, it was no persistent, lasting no longer than 7 days. Feeding regimen affects the timing of the clitellum development; yet regardless that feeding regime, all leeches exhibited two periods with a clearly shows clitellum. The very first appearance that a clitellum did not coincide with either spermatogenic or oogenic maturity; however at the second appearance that a clitellum, completely mature ova were present. The visibility of a visible clitellum is therefore not a good indicator the maturity in E. Obscura and also probably no a an excellent indicator in many or all types of Erpobdellidae. The visibility of a spermatophore is likewise not a an excellent indicator of maturity. Singhal et al. (1985) found that just 5% of maturation E. Obscura collected from the field had observable spermatophores, and also Biernacka and Davies (1995) found that only 4% of the mature-size-range animals had a spermatophore attached.

Erpobdellid cocoons are thick-walled, oval (Figure 23.10), and attached to a for sure substrate (stone, leaf, stem, wood). Cocoons that Barbronia weberi are comparable and are regularly attached to the pipeline of water plants (Govedich et al., 2003). The cocoons of Piscicolidae and Hirudiniformes are just loosely attached come the substrate and are typically spherical. Species of Hirudiniformes deposit your cocoons in moist habitats the end of water; the outer wall surface of the cocoon is spongy, i m sorry is believed to reduce water loss. The cocoons that Glossiphoniidae are very thin-walled and are one of two people deposited on the substrate and also immediately covered by the parent’s human body or attached to the ventral surface of the parental (Fig. 23.10). In both cases, the hatchlings space attached come the ventral body wall surface and are carried approximately by the parent for a long period. For Theromyzon tessulatum Müller, 1774, this can be 5 months; and also for Theromyzon trizonare, this deserve to be one month (Wilkialis and Davies, 1980a). Glossiphonia complanata adults administer their young through nutrients that are passed through the body wall surface of the parent into the posterior sucker the the young (de Eguileor et al., 1994). To reduce in human body weight and also size have actually been tape-recorded for brooding glossiphoniids; however, these losses have actually been meeting to reduced feeding potentials and to the energy expended ventilating the young (Calow and Riley, 1982; Milne and Calow, 1990). It is possible that other varieties that brood attached young may carry out nutrients come the emerging young; however, more studies have to be conducted on a variety of glossiphoniid leech species. A compare of reproductive output for glossiphoniids that brood eggs and young and also for erpobdellids the deposit egg in a cocoon mirrors that brooding is no metabolically more expensive than encapsulation. There space no extra energy prices incurred in terms of carrying or ventilating broods, however weight loss because of reduction in feeding occurs. This costs appear to be energetically similar to the expense of encapsulation for erpobdellids; and therefore, brooding and encapsulation represent alternate evolutionary courses for brood protection.

Globally, some 6000 earthworm types have been described, currently under a variety of climatic regimes native boreal to tropical biomes (Butt and also Lowe, 2011). Every earthworms have actually a similar life cycle. ~ a period of weeks come months, an earthworm cocoon will hatch in the floor or leaf litter to produce a hatchling worm. This pet is complimentary living and also grows over a duration of weeks or months to maturity, the latter shown by the presence of a clitellum (saddle). Earthworms room hermaphrodites (i.e. They act together both male and female), but most species reproduce sexually by exchange of sperm (amphimictic), whilst rather reproduce asexually (parthenogenetic). Cocoons are developed periodically through adults v secretions native the clitellum, and they room deposited in the soil (Fig. 20.1). The exact timescales and also numbers produced are species specific.

Adult sizeSmall-mediumMediumLarge
Reproductive rateHighIntermediateLow
PigmentationPredominantly red colorationPredominantly unpigmentedPredominantly brown to black coloration
MobilityHighly mobile—selected members the this group are considered pioneer speciesIntermediateLow mobility, generally found in undisturbed habitats, inhabiting semipermanent burrows
Location in the floor profileOrganic horizons in or near the surface ar litterUsually found in the soil profile inhabiting horizontal burrows within 15 cm that the surface (in pleasant locations)Construct semipermanent upright burrows (to a depth the 2 m) in the soil profile. Burrows open onto the floor surface
Food sourceDecomposing surface litterOrganic matter ingested through soil (geophagous)Surface litter attracted into the burrow—usually at night
Temperate examplesEisenia fetida, Dendrodrilus rubidusAporrectodea caliginosa, Aporrectodea roseaAporrectodea longa, Lumbricus terrestris
Tropical examplesPerionyx excavatus, Eudrilius eugeniaePontoscolex corethrurus, Millsonia anomalaAmynthas rodericensis, Eutyphoeus gammiei

Source: adapted from Butt and Lowe (2011).

For the purposes of this chapter, the is more relevant to separate earthworms into two distinct categories—those that inhabit organic issue (litter or compost dwelling) and also those the inhabit the mineral soil (soil dwelling).

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Tsuyoshi Kawada, in Handbook that Hormones, 2016


Structure and also Subtypes

Genomic evaluation of the annectocin receptor (AnR) gene revealed the the intron-inserted place is conserved between the AnR gene and also mammalian OXT/VP gene <3>. In situ hybridization using an earthworm showed that the AnR gene was particularly expressed in the nephridia situated in the clitellum region <3>. One more annelid receptor homolog for the OXT/VP suprafamily peptide to be cloned native the leech Theromyzon tessulatum <4>, return no endogenous ligand has been found in the leech. The receptor homolog gene is express in the genital tract, ovary, and mind of the leech. Furthermore, leech AnR mRNA slowly increases during the duration of sexual maturation, and also disappears after egg-laying <4>.

Signal Transduction Pathway

Administration that annetocin to Xenopus oocytes to express AnR induces Ca2+-dependent signal transduction. The EC50 value of annetocin for AnR to be calculated to be around 40 nM by dose–response curves for present shifts <3>.

Tarmo Timm, Patrick J. Martin, in Thorp and also Covich's Freshwater Invertebrates (Fourth Edition), 2015

Reproduction and Life History

The individual lifetime of tubificids and lumbriculids in aquaria often lasts numerous years and may incorporate several reproduction cycles. Some people of T. Tubifex and also Spirosperma ferox have actually reached an age of 10 or more years. The earliest recorded period was because that an separation, personal, instance of C. Lacuum (an aquatic “earthworm”), which live 46 years in an aquarium, although there is no reproducing. The actual life time of the same varieties in nature is undoubtedly shorter due come predation. The facultative parthenogenetic varieties T. Tubifex deserve to survive and also reproduce in aquaria for plenty of years, even at a persistent room temperature, as have the right to the architomic clones that the tubificids Bothrioneurum and also Aulodrilus. Some other varieties can live several years without reproduction uneven “hibernation” allows them to experience the common sexual procedure (Timm, 1987). Most types reproduce once a year and also then reabsorb the entirety system apparatus (except gonads) and also apparently end up being immature till the next sexual cycle. Oligochaetes room hermaphrodites, greatly protandric, with the partners trading sperm throughout copulation. The clitellum and also some one-of-a-kind cutaneous structures, occasionally in combination with penial chaetae (Cuadrado and Martínez-Ansemil, 2001; Caramelo and Martinez-Ansemil, 2012), assist to hold the partner together during this process. The alien sperm is save in spermathecae either together amorphous mass, organized into bundles of different complexity, or as spermatozeugmata consist of of 2 kinds of spermatozoa, the last characteristic the the subfamily Tubificinae (see Jamieson and Ferraguti, 2006). As soon as spermathecae are absent, the spermatophores are then attached to the exterior of the partner’s body; this wake up in the tubificids Bothrioneurum and Paranadrilus, the crassiclitellate Criodrilus, and also some others. Both the egg(s) and alien sperm will be laid right into a cocoon secreted by clitellum. The mother worm sheds the cocoon through crawling backwards, after i beg your pardon the cocoon’s shell will harden and also the ends will contract into a sealed plug. Eggs are fertilized within the cocoon, and development is direct. After some weeks, the young worms will leave the cocoon. “Microdrile” aquatic worms place large, yolk-rich eggs. In contrast, the eggs of “megadriles” room smaller, and also embryonic expansion depends ~ above nutritious fluid secreted into the cocoon by the mom clitellum. Spermatogenesis occurs previously than oogenesis and also at reasonably lower temperatures. This describes the observation that in a moderate, seasonal climate numerous aquatic oligochaetes mature and also copulate in winter or beforehand spring and lay egg in the beginning of the warmer period.

Although oligochaetes might reproduce only when per year in seasonally cool aquatic systems, some species (e.g., L. Hoffmeisteri) have the right to mature quickly in warmer water and also produce number of generations every year. Some types (e.g., T. Tubifex and also Ilyodrilus templetoni) can develop parthenogenetic eggs, as confirmed by absence of spermathecae in plenty of individuals. Part rare taxa likewise lack the masculine apparatus, as in Tubifex pomoricus. Parthenogenesis is a method to prevent the temperature trouble with spermatogenesis and to produce several generations a year, together in the instance of T. Tubifex and L. Hoffmeisteri. However, fecundity progressively decreases over several parthenogenetic generations.

Another kind of asexual reproduction that fully avoids the resource- and time-consuming sexual procedure and egg-laying is vegetative reproduction, either by architomy or paratomy. Architomy way enhanced capability to regenerate a complete pet from separate pieces, after either a violent assault or voluntary fragmentation. Architomy is almost obligatory in L. Variegatus however common likewise in part Tubificidae (e.g., Bothrioneurum, Aulodrilus, Potamothrix bedoti) and Enchytraeidae (Cognettia and some others). At the very least in these tubificids, architomy is characteristics of warmer periods, whereas at reduced temperatures, the procedure ceases and at least some component of the populace becomes sexually mature. In Lamprodrilus mrazeki (Lumbriculidae), architomy wake up in cysts created in summertime once pools dried up.

A an ext advanced technique of asexual reproduction is paratomy: the production of a chain of zooids by longitudinal budding. This characterizes entirety families that Naididae, Pristinidae, and also Opistocystidae. Paratomy deserve to prevail throughout favorable seasons, when sexually mature people appear and also produce cocoons mostly prior to a colder or dryness period. The parental then generally die after forming cocoons.

M.E. Benbow, in Encyclopedia the Inland Waters, 2009


The phylum Annelida is stood for by 5 classes of freshwater worms, through the Oligochaeta, Polychaeta, and also Hirudinea being the most usual groups encountered in inland freshwater systems. The continuing to be classes include the Branchiobdellida and also the an extremely rare Archiannelida, the former virtually exclusively represented by commensals or parasites of crayfish. Much of this rundown of the freshwater Oligochaeta and also Polychaeta is synthesized indigenous references listed in the further Reading section.

The Annelida comprises the segmented worms, including earthworms (Oligochaeta), leeches (Hirudinea), and the generally marine tubeworms (Polychaeta). The basic annelid human body is a linear collection of virtually identical segments, other than for the prostomium (head) that residences the brain, and the pygidium, i beg your pardon is the terminal part of the body the holds the anus. Number of burrowing and also tube-dwelling polychaete taxa have actually modified components of the body that are distinctive regions of segments. Annelids usage coelomic fluid pressure together a hydraulic skeleton, which functions as one antagonist v longitudinal muscles in a peristaltic manner, to move the organism front along peak of and within substrata. Annelids are hermaphroditic (having both male and also female reproductive organs) and the freshwater representatives room usually syngamic (i.e., organisms through gamete fusion) through cross-fertilization; some varieties are known to reproduce by budding. The Oligochaeta and also Polychaeta are morphologically distinct and also easy come distinguish, however both are typical to the depositional area of lakes, wetlands, streams, and also rivers wherein they assume a decompositional function in organic issue processing. The these 2 understudied class of freshwater annelids, the oligochaetes are much better known due to the fact that they are relatively much more diverse and also abundant throughout temperate zone areas of the world, if polychaetes are reported come be more widespread throughout the tropics, with only a couple of species reported from warm latitudes.


There room at least 10 family members of oligochaetes that have actually freshwater representatives. Four households are strictly aquatic and include the Aeolosomatidae, Naididae, Tubificidae, and Lumbriculidae. There space several other taxa from various families that are taken into consideration amphibious or semiaquatic since of accidental wash-ins during flooding or other disturbances. Although substantial studies are lacking for the Oligochaeta, most taxa (e.g., Tubifex) space thought to have a cosmopolitan circulation with a couple of representatives minimal to particular regions of the civilization (e.g., plenty of Lumbriculidae). A unique types of the Enchytraeidae (Mesenchytraeus solifugus – ice worms) is discovered only ~ above the glacial ice cream sheets that the Pacific Northwest, where populaces feed virtually exclusively on tree pollen and also algae.

Oligochates display distinct metamerism, whereby the human body is composed of virtually identical human body segments serially aligned follow me the length of the body. The number of segments varies by species with some taxa (e.g., Haplotaxidae) with virtually 500 segments, however most taxa through usually less than 200 (e.g., many tubificids). Aquatic oligochaetes are typically smaller 보다 their terrestrial relatives, hardly ever reaching body lengths greater than 30 mm. On each segment are discovered setae, small, stubborn hair-like structures that offer as exterior grips because that locomotion. In general, aquatic types have much longer setae than carry out their terrestrial relatives. The setae room extended during longitudinal muscle contractions for movements that are assisted in by peristaltic contractions of the coelomic (body cavity) fluid. The direction the the body wall contractions can be reversed to enable for behind locomotion. Many aquatic oligochaeta relocate over or within the interstitial spaces that substrata, yet a couple of are well-known to be reasonable swimmers (e.g., some types of Naididae) that relocate in a serpentine way immediately over the substrate.

Gas exchange takes location through the slim body wall surface in all aquatic oligochaetes; however, some tubificids and also naids are known to pump water right into the anus to effectively increase the body wall surface surface area easily accessible for gas exchange, and other taxa have ciliated anal gills that offer a supplementary duty to body wall gas exchange. In addition, some species of tubificids additionally have respiratory tract pigments that help in respiration. These added respiratory mechanisms enable those taxa come survive and also occupy habitats with an extremely low liquified oxygen.

Although absent in all various other oligochaetes, many varieties of Naididae have epidermal eyespots through photosensitive receptors. The evolutionary meaning of these frameworks within a single family that oligochaetes is tho debated, as numerous polychaetes and a few Hirudinea have comparable structures. Oligochaetes additionally possess additional sensory structures that are known to aid in tactile, thermal, and also chemical detection.

Oligochaeta are commonly hermaphroditic and most have actually sexual reproduction that involves reciprocal sperm transfer and fertilization that takes ar in a cocoon that is secreted through the clitellum; however, number of taxa the the Naididae and Aeolosomatidae give birth asexually via budding. The clitellum is a reproductive part of the oligochaete body consisting of collection adjacent segments whereby the epidermis is swollen and also house glands that secrete mucus for copulation. In many sexually reproducing aquatic taxa, the clitellum is just two segments broad and sometimes only one cabinet thick making it difficult, if not impossible, come detect with the nude eye. For part terrestrial taxa the clitellum can be as an excellent as 60 segments wide and plenty of cells thick, offering it a ‘girdle’ appearance. The mucus from the clitellum creates a chiton-like cocoon framework where fertilization and also embryonic breakthrough occurs. This cocoon is believed to play a function in desiccation resistance and also dispersal abilities of aquatic oligochaeta.

Aquatic oligochaetes occupy the depositional habitats of many aquatic habitats, functioning together decomposers the decaying necessary matter and also mixing and also aerating the benthic substrates with burrowing. Density of aquatic oligochaete is typically low in streams, however can it is in appreciably high in eutrophic habitats through low liquified oxygen or anoxic conditions. Indeed, aquatic oligochaetes room often taken into consideration indicators the nutrient-enriched flowing waters as soon as in relatively high abundance contrasted with other benthic invertebrate taxa. Plenty of aquatic oligochaetes have the right to withstand lengthy periods that time under anoxic, or near anoxic, conditions. Because of this, oligochaetes are frequently most plentiful in wetlands and marshes and can achieve big benthic densities in the deep area of lakes. For instance, populaces of some tubificid worms can accomplish densities as good as 8000 people per square meter. However, there have actually not been considerable studies pertained to the ecology of details aquatic oligochaetes beyond wide taxonomic surveys or studies where oligochaetes were gathered as part of much more general benthic invertebrate research.


Of 85 family members in the course Polychaeta, only 10 have representatives in brackish and freshwater habitats. Over there are an ext than 5000 identified species in this class, with much less than 2% classified as euryhaline or freshwater and also less than 50 species minimal to freshwaters. Most freshwater and euryhaline representatives are within the Nereididae (Figures 1 and 2), which is predominantly a naval family. However, there are other families containing a couple of euryhaline and freshwater types (i.e., Phyllodocidae, Nephtyidae, Spionidae, Sabellidae, Serpulidae, and Ampharetidae). Over half of the 50 euryhaline and freshwater species of Nereididae are located in the tropical and also subtropical western Pacific region, while only 10 species are discovered in north America. Freshwater and also euryhaline polychaeta have actually been report from phibìc America, including Alaska, California, Georgia, the great Lakes, brand-new York, caves the Mexico, and also the Philippines, Canada, Papua brand-new Guinea, Fiji, brand-new Zealand, the Amazon, Japan, China, India, and Hawaii, and also probably other regions that the world. Many freshwater or brackish polychaete habitats were when geographically linked or room presently linked to the ocean. In addition, many specimens have been gathered within about 30 km the the ocean, supporting a concept of recent evolution into freshwater. Interestingly, it has actually been reported that part freshwater taxa show direct breakthrough rather 보다 the an ext usual larval advancement of marine representatives, a feature similar to the freshwater family members of a couple of other predominately naval invertebrates (e.g., crabs, shrimps, bivalves).



The general polychaete body setup is unique from oligochaetes in that each segment is defined by two lateral muscular appendage-like structures called parapodia (Figures 1 and also 2). Every parapodium consists of setae (usually multiple) that are offered in locomotion that can take place along the surface of substrates, in ~ the interstitial spaces that substrates, or for swimming, while part are progressed for a tube-dwelling existence; however, this latter life format is no common amongst the freshwater forms. Most polychaetes have actually a highly emerged prostomium that includes sensory guts that include antennae and also eyespots (Figure 2). Plenty of of the nereid taxa have actually pharyngeal jaw-like structures, located several segment posterior of the prostomium, the are provided for catching prey or shredding detritus (they can additionally leave a nasty bite if provoked when handling).

Very small is known about the ecology that freshwater polychaetes. The species that have actually been studied many are the Nereididae, and also are many often gathered from extremely organic depositional habitats of streams, wetlands (and rice paddies), and also lakes. A couple of species have actually been documented to have actually an arboreal existence in phytotelmata (e.g., little collections the rainwater linked with plant sheet axils or in bromeliads), but little is known about their ecology. Choose their oligochaete relatives, polychaetes appear to be rather tolerant to low dissolved oxygen conditions. The Namalycastis species often have actually a hemoglobin sinus the runs the length of the body, and also it is presume to contribute to respiration in anoxic problems of depositional habitats (Figures 1 and also 2). In tropical areas of the world, freshwater polychaetes are frequently abundant in the greatly polluted get of streams and rivers, possibly making them bioindicators that anthropogenic nutrient enrichment. The benthic polychaete taxa are thought to be scavengers and also detritovores; however, that is most likely that some nereid creates are energetic predators as well. Most freshwater polychaetes space thought to ingest any and all debris, assimilating the connected microbial flora and also organic matter, and they might play a notable duty in decomposition in sluggish flowing dry streams. Very small is recognized of the reproductive strategies of freshwater polychaetes, but most literature suggests that they have the right to reproduce sexually or asexually, and also then go v direct development with no larval stage.

One of the relatively more well-documented nonnereid species of freshwater polychaete is Manayunkia speciosa (Sabellidae) that has been report from plenty of locations in north America, consisting of the good Lakes region, brand-new Jersey, Georgia, brand-new York, California, Alaska, north Carolina, and South Carolina. This types is generally less 보다 5 mm in body length and also secretes mucus to form a mud/silt/sand pipe in i beg your pardon it resides in a filter-feeding fashion, utilizing ciliated tentacles the the prostomium. In temperate regions of the world, this is more than likely the many common types collected by benthic ecologists.

Annelids blee both sexually and asexually. Sexual reproduction is most common amongst all main families of Oligochaeta and also Hirudinea. Throughout copulation, two mature, hermaphroditic animals embrace in such a way that your heads face opposite directions. In this position the male genital pores room opposite to the spermathecal opening of the various other (Figure 20.8). Copulatory structures, amendment musculature of the body and also genital chaetae, aid in the exchange the spermatozoa. In some families, accessory glands create a secretion that ‘glues’ partners together. Copulation last around 5–10 min. The spermatozoa are stored in the spermatheca till the cocoons room laid. Cocoons are created by a clitellar gland secretion developed in the clitellum the acts as cocoon membrane. The membrane initially is elastic and transparent. By contractions that the body wall musculature, the membrane slides towards the head. As soon as passing the mrs genital pore, the eggs room pushed into it; and when happen the spermathecal opening, the spermatozoa room added. The fully formed cocoon is deposited in the upper layer the the substrate or fastened to tough material, like macrophytes. After ~ the cocoon is deposited, that hardens once the membrane becomes chitinous and takes a distinctive shape. Cocoons can include up to 40 eggs. The cocoons space deposited a considerable time period after copulation due to the fact that the mrs reproductive device takes much longer to develop. Cross-fertilization is usual in annelids, however self-fertilization may also occur. The timing of sex-related reproduction varies widely among annelid types in an answer to various environmental conditions. Some develop cocoons throughout the year, whereas others are restricted to certain weeks.

In naidids, asexual reproduction (mostly by paratomy) is considered the dominant type, while sex-related reproduction wake up rarely and also then only sporadically. Beforehand in the twentieth century, Piguet (1906) observed that sex-related reproduction the some varieties occurred seasonally in certain sites. To survive periods of unfavorable conditions, especially droughts, protective cocoons allow populations to withstand stress and recover once favorable environmental conditions return. The cocoon is an reliable trait adaptation to make it through adverse conditions. In this regard, the cocoons the naidids in excessive habitats space analogous to the gemmules that sponges, the relaxing eggs that rotifers, and the ephippia of cladocerans (Parish, 1981). Christensen (1984) verified that a good variety the asexual reproductive settings are known amongst aquatic oligochaetes. He distinguished four main varieties of asexual modes and indicated the genetic and ecological implications, as follows. First, the often expressed expectations of a strong predominance that one specifically adaptive genotype space not born out. Second, in most cases, a variety of genetically unique clones are present in each population, and they present a solid differential distribution in heterogeneous environments, denote an reliable exploitation the the accessible resources. Third, most instances of asexual propagation room reproductive strategies of your own and not escape mechanisms. And finally, the mechanisms underlying asexual propagation are complex and show off many facets of the life history.

The ability of tubificids to reproduce parthenogenetically following, or in place of, bisexual reproduction has been displayed (Poddubnaya, 1984). Throughout parthenogenesis, the spermatogenesis ceases at very early stage of sexual cell development. Together with some peculiarities of the structure of the sex-related system, this rules the end the possibility of self-fertilization in this household of worms.

Nikolai J. Tatarnic, in recommendation Module in Life Sciences, 2018

Male-Male Competition

The evolution of copulatory wounding as a viable mating strategy is partly described as result of competition among males, together they vie come fertilize as many female eggs as they can, come the exemption of their competitors. In the bean weevil Callosobruchus maculatus copulatory wounds inflicted by spines top top the masculine genitalia damage the mrs genital street (TP) (Crudgington and also Siva-Jothy, 2000) to reduce female lifetime reproductive success (Rönn and Hotzy, 2012). Males with longer spines accomplish greater adjustment success in sperm compete (Hotzy and also Arnqvist, 2009; Hotzy et al., 2012). Interestingly, rather than reducing female willingness to mate, genital damages apparently leader to the opposite, with females remating sooner and laying under eggs, potentially limiting the services to males. Wounding per se may thus be a pleitropic by-product of choice on various other reproductive characteristics that rise male fitness (Morrow et al., 2003). In the simultaneously hermaphroditic earthworm Lumbricus terrestris, wounds caused by copulatory setae situated ventrally on the body, far from the clitellum (the worm’s reproductive organ), enable for the move of secretions that induce raised sperm absorb (TST) (Koene et al., 2005). Keep in mind that in both instances wounding is no under direct selection: in the first it is likely a pleitropic side-effect, and in the 2nd the wounds are just a mechanism through which antagonistic secretions are transferred.

Evidence for the underlying function of sperm vain in control TI in part systems deserve to be uncovered in the mating system of the traumatically inseminating spider Harpactea sadistica (Řezáč, 2009). In most spiders, males inseminate the female with her epigynum (genital opening), and the sperm then pass into her spermatheca, a sperm storage body organ with a solitary opening. As is usual in numerous arthropods, fertilization occurs just before oviposition: the egg descend right into the uterus, where sperm is then released native the spermatheca, and also the woman lays freshly fertilized, single-celled eggs. Where females possess a cul-de-sac kind spermatheca through one opening, either the sperm the the last male to mate through the mrs are offered first, together they reside closest to the spermathecal opening, or rather the sperm from multiple matings i do not care mixed, calculation paternity unpredictable. In H. Sadistica, TI enables the masculine sperm to bypass the spermatheca (which is atrophied in this species), and also instead travel directly to the ovaries where fertilization occurs, with females eventually laying partially occurred embryos. In this instance TI is believed to enable males come “escape” sperm competition by alternating the typical order of sperm precedence (Řezáč, 2009). In traumatically inseminating Cimicomorpha (bed bugs and also their allies), fertilization likewise occurs in the ovaries, prior to ovulation. However, ovarian fertilization wake up in several various other cimicomorphan lineages as well (e.g., the household Miridae (Wheeler, 2001), which has a single TI genus, and the Tingidae (Marchini et al., 2010), which have actually no recognized TI species), indicating that ovarian fertilization can be accomplished without TI, and thus might not be result of TI in the Heteroptera. It is clear the there is lot to learn around reproduction in this taxa before we deserve to draw any kind of firm conclusions ~ above the relationship between TI, sperm storage, and fertilization.


Laboratory name and address:

Laboratorio de Ecologia de Agroquímicos, Instituto Biológico, São Paulo SP, Brazil


Contact persons:


Title of the analytical method:

Estimation of the bioaccumulation element of radiolabeled pollutants in compost worms Eisenia andrei/fetida



As earthworms room an integral component of numerous soil food webs, they might serve as bioindicators because that the potential dangers posed by pollutants (pesticides, drugs, and metals) to soil biota. The an approach presented is a activities bioassay to evaluate the bioaccumulation that radiolabeled (using a 14C-label) pollutants in compost worms that prosper in soil and also incorporate a specific amount of radiolabeled pollutant. Fluid scintillation counting is offered to estimate the amount of radioactivity current in essential extracts all set from compost worms and soil.



The current technique is applicable come the determination of radiolabeled pollutants in soil utilizing earthworms as soil bioindicator organisms.


Equipment and instruments:•Liquid scintillation respond to (LSC)

Laboratory microwave

Rotary evaporator




Reagents and also materials:•Organic solvents, residue evaluation grade

14C-labeled target pollutant

Pollutant (nonlabeled)

Adult Eisenia andrei earthworms


Detailed procedure (protocol):•One week before the start of the study, sweet 200 g samples of different soils in 500 mL glass beakers and also moisten come 60% that the best water holding capacity (MWHC) v deionized water, together recommended by OECD overview Nº 317.

Weigh sufficient sample replicates to have at the very least three floor replicates for the treatment with the pollutant and also three replicates for the control.

For acclimation to laboratory conditions at the very least 1 day prior to the start of the study, place five adult earthworms every replicate each weighing an ext than 300 mg with clitellum in untreated soil with at the very least 50 g floor per earthworm.

On day 1 the the study, slowly treat and mix the soils with a systems of 14C-target pollutant. The samples stay in a fume hood until the solvent has evaporated.

Only after the solvent has evaporated native the soil the acclimatized adult earthworms are placed in the cure or control soil.

Cover the surface ar of the glass beakers through perforated plastic movie to enable air exchange.

Weigh each complete glass maker containing the moistened soil and also the compost worms to evaluate the dynamics that water evaporation throughout the study duration (Vampré et al., 2010).

Keep the glass beakers in ~ a continuous controlled temperature (e.g., 25°C) and standardized light durations (e.g., 12 h).

Every 3 days sweet the beakers and adjust the water content to keep the floor moisture in ~ 60% MWHC.

After 14 days, disassemble the beakers and separate the worms; wash in madness water, dried in filter file and weigh to compare v the initial weight and determine the result of the pollutant on expansion of the organisms.

Maintain the earthworms in ~ 4°C because that 24 h ~ above wet filter paper, i m sorry is adjusted after two durations of 12 h to allow emptying of floor or substrate particles from your guts. Freeze because that 24 h. Monitor the extraction technique by Papini et al. (2006) to recuperate the pollutants from the necessary extracts ready from compost worms and soil. All organisms of every replicate room extracted with each other by this method.



The bioaccumulation variable is established by splitting the lot of pollutant current in the earthworms (Co, in kBq/g or µCi/g) by the quantity in the soil (Cs, in kBq/g of soil dry weight):


Quality control:

Preliminary experiments should be carried out to recognize the extraction recovery the the target pollutant from the soil and earthworm tissue. The recovery have to be at least 85% for the added 14C-labeled target pollutant.

To calculation the extraction recovery indigenous earthworms, wash 5 adult worms (with clitellum and much more than 300 mg each) in madness water in a plastic sieve, dry closely in filter file and freeze at −18°C because that 24 h. Spike the frozen animals with a solution containing a mixture that the 14C-pollutant and also a reasonable—but not lethal—amount of the pollutant. Cut the animals into small pieces of around 1 cm and also homogenize for about 15 min under a fume hood to allow evaporation that the spiking solvent.

Material native the cure earthworms is extract by enhancement of a solvent or a solvent mixture by microwave-assisted exploit (MASE) making use of frequent quick (a few seconds) low power cycles as the earthworms save on computer a far-ranging amount the water. The heating right into microwave is intercalated through temperature decrease with immersion in ice-bath (method by Andréa et al., 2001). Relying on the solvent used, few mL aliquots of the extract obtained are directly counted through LSC making use of a scintillation cocktail. If the solvent is a quencher, it deserve to be rotary-evaporated and also resuspended in another an ideal solvent compatible through liquid scintillation counting.

To estimate the recovery following extraction native soil, an adequate soil extraction an approach should be established. For example, the samples might be extract by MASE in assorted cycles of few seconds and low level of microwave energy, also intercalated through ice-bath. The extraction recovery should be established in triplicates utilizing 3.0 g wet samples spiked with a recognized amount of the pollutant.

Extraction recovery is calculated together the ratio between the amount of 14C-pollutant recovered in soil and also earthworm extracts, and the lot of 14C-pollutant added into the samples, taking into account the dilution factors.



For a check to it is in valid, the mortality that the bioindicator organism must not exceed 10% of the total number of earthworms in every replicate of control or cure samples.

Ensure the the measurement conditions of the LSC space checked to permit valid measurements. These encompass daily check of the carbon-14 calibration standard and also the typical of lift radiation to verify the efficiency of counting.


Interferences, troubleshooting, and also safety:

All laboratory personnel need to be trained on the safety and also correct handling of the pollutants, radioactive materials and also the quenchers for LSC.



Andréa, M.M., Papini, S., Nakagawa, L.E., 2001. Optimizing microwave-assisted solvent exploit (MASE) of pesticide from soil. J. Environ. Sci. Health. Component B, Pesticides, Food Cont. Agric. Wastes, B 36, 87–93.

OECD – (Organization for economic Co-operation and also Development), 2010. OECD guideline Nº 317. Bioaccumulation in Terrestrial Oligochaetes. 30 pp.

Papini, S., Langenbach, T., Luchini, L.C., Andréa, M.M., 2006. 14C-Paraquat in compost worms Eisenia foetida. J. Environ. Sci. Health. Part B, Pesticides, Food Cont. Agric. Wastes, B 41, 523–530.

Vampré, T.M., Fuccillo, R., Andréa, M.M., 2010. Oligoqueta Eisenia andrei como bioindicador de contaminação de solo por hexaclorobenzeno. R. Bras. Ci. Solo 34, 59–66.


Minimum method validation data:

The technique validation was performed using 14C-Paraquat and 14C-Simazine herbicides (Tables 3.36 and 3.37).

Table 3.36. Radiocarbon recoveries indigenous 5 (five) earthworms spiked v 14C-herbicides using different methods

MethodSolventVolume (mL)ExtractionConditionsRecovery (%±SD)
1Ethyl acetate100Shaker4 cycles, 6 h each100.2±2
2Methanol150Shaker5 cycles, 6 h each88.9±2
3Methanol150Soxhlet1 cycle, 8 h116.8±5
4Methanol:dichloromethane (10:5)15Microwave15 cycles, 30 s each and also 1120W99.2±8

MethodSolventVolume (mL)ExtractionConditionsRecovery (% ± SD)
1Methanol100Shaker3 cycles, 8 h each97.1±2
2Methanol100Soxhlet1 cycle, 12 h105.1±2
3Water:methanol:dichloromethane (1:8:6)15Microwave10 cycles, 20 s each and also 800W89.4±1
4Hexane:Acetone (75:25)15Microwave12 cycles. 2 min each and also 480W81. 1±1

The percentage full radiocarbon recovery together 14C-extractable + 14C-bound in soil and earthworms in ~ the finish of the experiment was: 93.9±9%.

The selection of sports on the performance of the organic combustion an approach to determine the 14C-bound in soil and earthworms to be 91%–95%.

The limit of Detection (LOD) was calculated as the typical + 3 time of the task (dpm) typical deviation the the blanks (6:4 of monoethanolamine and scintillation cocktail), estimated as 285 dpm (4.8 Bq).

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The border of Quantification (LOQ) was defined as the median + 10 time of the activity (dpm) typical deviation that the blanks, estimated as 593.0 dpm (9.9 Bq).