Neri, Floramae Joyce BS Bio 3 Bio 4.

1 YA- Invertebrate Zoology

Tabulated Form of the Classes Under Phylum Annelida Phylum Annelida Segmentation annelid body composed of 3 regions: Form and Function Classes Under Phylum Annelida Class Polychaeta Class Oligochaeta Class Hirudinomorpha body is dorsoventrally similar to that of annelids with legs flattened and tapered at the generalized annelids anterior end has well developed segments, 4 bundles of parapodia- legs; lateral chaetae per segment, small segments at both fleshy outgrowth of the body prostomium lacking extremities have been wall appendages, small modified to form suckers pygidium also devoid or appendages chaetae- simple, terminate anterior sucker- surrounds segment bears one pair of in a needle like point or tips the mouth; smaller than the parapodia that are bifid, pectinate posterior biramous consisting of: chaetae on genital segments are more complex chaetal sacs- located on each side of segments; chatae are secreted and which they emerge as bundles 2 chaetae bundles are ventral and two are ventrolateral or dorsolateral number of chaetae per segment varies from 1 to 25 posterior sucker- disc shaped and turned ventrally segmentation very much reduced; fixed number of 33 segments

*prostomium- contains brain and sense organs; anterior end of trunk

*trunk *pygidium- posterior end of body;ludes the anus segments- longitudinal series of similar body units peristomium- ventrally surrounds the mouth; behind the prostomium prostomium and pygidium are not considered as segments growth zone- localized to a region immediately in front of the pygidium

*notopodium- upper division

*neuropodium- ventral division parapodial muscles- attach to the acicula; move the parapodia cirri- project from the dorsal base of the notopodium and from the ventral base of neuropodium chaetal sacs- secretes a bundle of chaetae chaetae- used to improve traction for locomotion

chaetae are absent head consists or a reduced prostomium plus 5 segments head bears ocelli and is ventrally modified to form the anterior sucker trunk is consist of 21 segments; encompases the preclitellar region, clitellum, posclitellar region ventral, posterior sucker is derived from 3 segments

cells in growth zone: *paired ectodermal cells

*mesodermal teloblast cells teloblastic growth- body growth; succesive addition of segments posteriorly all internal structures are segmented chief integrating structures: *nervous system *hemal system *musculature *digestive system

some chaetae are used for: *spatulate shovels used for digging bear velcro-like hooks used to grip the inner walls of tubes and burrows paddles used for swimming having several different types of sensory appendages (unique to annelids) peristiomium bears: *sensory tentacular cirri *tentacular palps- 2 long feeding appendages prostomial appendages includes anterior/anterodorsal antennae and anteroventral palps

dorsal anus is on or near the last trunk segment in front of sucker

segmental and repeated in each segment: *appendages *coelomic cavity *nephridia Body Wall consist of: *fibrous collagenous cuticletoughen the body wall, resist bulges, impart iridescent sheen to the body

similar to annelids in general tubes- unique to annelids; may be open at one or both ends and partially buried or attached to surfaces

a thin cuticle overlies an epidermal layer that contains mucus gland cells circular muscles are outermost and the well developed longitudinal muscles are in 4 bands

consist of typical annelidan cuticle and epidermis fibrous connective tissue beneath epidermis is very thick and occupies much of the anterior body below the dermis is a layer of circular musculature followed by diagonal muscles and a powerful longitudinal musculature dorsoventral muscles present

*glandular monolayer epidermis *connective tissue dermis/cutis

function of tubes:

setae are relatively complete each coelemic compartment is connected to the outside

*protective

*musculature derived from coelomic lining

*a lair from which to emerge and catch prey

coelomopore- situated in the intersegmental furrows and provided with a sphincter; aids in keeping the integument moist and may deter predator

musculature and other tissues occupy the expanded connective tissue region

mucus-secreting gland cellscommon in epidermis chaetae- chitinous bristles that project outward from epidermis to provide traction; arises from pit like epidermal follicle follicle cells- form the follicle wall chaetal base- anchored by hemidesmosomes to follicle cells obliquely striated musculature- arranged in outer circular and inner longitudinal muscles Nervous System anterior dorsal brain at the prostomium pair of ventral longitudinal nerve cords

*may provide a source of clean, oxygenated water *attachment of tube enables them to inhabit hard, bare surfaces *brood their eggs and young in the tube tube material- a fibrous protein that has the appearance and texture of cellophane, parchmet or silk

brain and ladder like nerve cords like those described in generalized annelid brain is large and lobed if head bears sense organs pedal ganglia- associated with segmental nerve cods at bases of parapodia; unique to polychates; important centers that control complex parapodial movements ventral nerve cord may be paired and ladderlike chief sense organs:

2 nerve cord fuse in the ventral midline and situated under muscle layers of body wall brain shifted posteriorly ventral nerve cord has: 5 giant axons, 3 large diameter axons in dorsal part of cord and 2 small diameter axons mid-dorsal axon- fired by sensory input from the head two dorsolateral fiber- fired with sensory input from posterior end of the body

similar to that of generalized annelids anterior and posterior ganglia are concentrated into masses brain located at the 5th segment; consists of the subpharengeal ganglia

brain- dorsal subpharyngeal ganglia

ladder like nervous system nervous sytem of few annelids is intraepidermal/ submerged in the coonective tissue below the epidermis

first 4 pairs of ganglia are fused to form a subpharyngeal ganglion ventral nerve cod consist of 21 pairs of free ganglia that form the nerve cord of the trunk and 7 additional pairs of ganglia that fuse to form the caudal ganglion associated w/ posterior sucker

large diameter giant axonsin longitudinal nerve cords; allows rapid impulse conduction and shorter response time giant axon is severed, locomotion is unaffected but escape response is blocked

*nuchal organs- pair of ciliated sensory pits or slits situated posterolaterally on the prostomium; chemoreceptive organ; eversible; important for detecting food *ocelli- found in surface of prostomium in 2, 3 or 4 pairs; best developed in errant polychaetes *statocyst- predominate among sedentary burrowers and tube dwellers; if destroyed, compensating ability is lost *some may also have antennae, palps and cirri prostomial ocelli- pigment cups; walls are composed of rod-like photoreceptors, pigment cells and supporting cells sensory appendages of the prostomium (antennae, palps), peristomium (tentacular cirri), parapodia (dorsal and ventral cirri) & pygidium (pygidial cirri) bear sensory cells

subpharengeal ganglionprincipal center for motor control and vital reflexes and dominates the succeeding ganglia in chain

chromatophores- under neural control; dramatic change in color due to pigment movement on these specialized cells specialized organs are2 to 10 pigment cup-ocelli and sensory papillae sensory papillae- small, projecting disc arranged in complete ring around 1 annulation

lack eyes

sensory structures:

have simple pigment-ocelli cups integument- well supplied with dispersed unicellular photoreceptors situated in the inner part of epidermis clusters of sensory cells form a projecting tubercle w/ sensory processes extending above the cuticle; appear to be chemoreceptor tubercles- form 3 rings around each segment; numerous in more anterior segment especially the prostomium

*unicellular receptors (photo/chemo/mechanorece ptors)

*ocelli and eyes

*statocyst

Coelom and Hemal

*chemosensory nuchal organs houses a bilateral pair of coelomic cavities isolated from neighboring segment by septa

gas exchange occurs across the general body wall; specialized gills are present

similar to described hemal system of generalized annelids

reduction of the coelom and septa

each septum is composed of:

specialized gills- thin walled, delicate; unprotected outgrowth of the body surface; not enclosed in gill chambers gills associated w/ the parapodia

have integumental capillaries hearts- expanded muscular regions of circumenteric vessels that link the ventral and dorsal longitudinal vessels; have valve in the form of folds in the wall gas exchange takes place by diffusion of gases through the body wall mucus-gland secretion and fluid discharge through the coelomopores facilitate gas exchange in earthworms specialized gills occur in few oligochaetes

lack paired segmental coeloms

*2 layers of mesothelium

coelomic cavity is continuous and uninterrupted

*a layer of connective tissue mesentery- formed by contact between the left and right coelomic cavities of one segment modified epitheliomuscular cell- form the body wall muscle contractile mesothelial cellmay form musculature on the septal faces, around blood vessels and around the gut chlorogogen cells- form a yellow or brown layer of tissue around part of the gut wall and certain blood vessels; plays a vital role in metabolism; center of glycogen and fat storage other function of chlorogogen cells: System

elytra- roof of overlapping fish-like scales and borne on a stalk gill ventilation- result from ciliary beating, muscular contraction of the gill or both circulation result from fluid movement in both hemal system and coelom parapodial vesselstransport blood to parapodia, body wall and nephridia before returning to dorsal vessel intestinal vessel- deliver blood from gut to ventral vessel

coelom is the global circulatory system replacing the hemal system two main longitudinal vessel are lateral and the vessels are lined by mesothelium coelomic mesothelium is specialized as large nutrient storage cells called chlorogen tissues coelomic fluid propelled by muscular contractions

larger oligochaetes have hemoglobin dissolved in blood plasma

aquatic ologochates tolerate low oxygen levels and for a short period of time, even complete lack of oxygen

general body surface providing for gas exchange

capillary plexus- found on the parapodia and gut wall; facilitates exchange in these organs gills- provided with afferent and efferent vascular loops permitting a two way flow blood- colorless in small polychaetes; some species large, extracellular molecules of hemoglobin

piscicolid gills- lateral leaflike or branching outgrowths of the body wall circulatory pigmentextracellular hemoglobin found only in arhynchobdellin leeches; responsible for the oxygen transport

*storage and detoxification of toxins *hemoglobin synthesis

occur in plasma *protein catabolism and formation of ammonia hemoglobin- most common respiratory pigment chlorocrourin- kind of hemoglobin; binds an atom of iron at its center; it is a non-heme protein; makes blood green hemerythrin- has pink or violet hue coelomic fluid- circulated by muscle contractions of body wall/by both muscle contraction and cilia in coelomic lining hemal vessel- hemoglobin binds less tightly muscle and nerve hemoglobin- binds it more tightly coelomic fluid hemoglobinbinds intermidiately hemal system hemoglobinbinds oxygen at gills mitochondrial cytochromeshave the highest affinity of all, final destination of respiratory oxygen torpid- rely on stored oxygen bound to hemoglobin

*synthesis of urea

coelomic fluid- circulated by cilia and contraction of body wall muscles; internal defense, gas transport coelom- fluid skeleton against w/c the muscles act to change body shape contraction of longitudinal muscles- body widens contraction of circular muscles- body elongation coelomic cavities- local segmental circulation well developed hemal system is responsible for transport throughout the body blood vessels and sinusessimple fluid channels in connective tissue compartment that lack vertebrate-like endothelium principal vessels: *dorsal blood vessel- dorsal mesentery *ventral blood vesselventral mesentery major blood vesselcontractile and pump blood peristaltically

hemoglobin- occur in coelomic fluid, blood, muscle, nerve; packaged in coelomocytes; dissolved in plasma of blood gas exchange occurs in: *body wall *appendages *gills excretory organs ore segmental nephridia metanephridial system occurs in annelids that have a hemal system, some vessels of which contain podocyte-covered filtration sites protonephridia are present in annelids that lack hemal system

Excretory System

filtration nephridia

have a metanephridial system one pair of metanephridial tubules per segment except t the extreme anterior and posterior ends holonephridia- a single typical pair of metanephridia typical or modified nephridia open to the outside via nephridiopores enteronephricnephridiopores open into various parts of the gut enteronephridia- restricted to certain parts of the body; adaptation for the retention of water excrete urea ammonia remains an important excretory product nephridia- regulates salt and water balance urine in both terrestrial and freshwater species is hypostomic

consist of 10 to 17 pairs of metanaphridia

intestinal walls- may play accessory roles in excretion

1 pair per segment

protonephridia occur in a few adult that lack or have reduced hemal system most have hemal and metanephridial systems metanephridial systemshave vascular filtration sites covered with podocytes and septum-associated metanephridia nephridiopore- opens at the base of the neuropodium on the ventral side; entire lining is ciliated metanephridia may be restricted to excretory segments protonephridium- bears a cluster of terminal cells solenocyte- terminal cell; resemble a sponge choanocyte, consisting of single flagellum in a collar of microvilli

nephridial tubules are embedded in connective tissue nephrostomes project into the coelomic channels mesothelium of the coelomic channels in the region of nephrostomes and canculi is porous nephridial tubule consists of a main duct that receives numerous ductules main duct expands into a urinary bladder before opening to the exterior at a ventrolateral naphridiopore

integument-where water absorption and loss occur capable of encystment during unfavorable environmental condition Digestive system and Nutrition straight tube that extends between the mouth and pygidial anus gut- passes through the septa and is supported above and below by mesenteries gut tube consist of: similar to the digestive system of generalized annelids annelids a straight tube extending from the mouth to the anus scavengers that feed on dead organic matter they also deposit feed; ingest organic matter obtained from mud or soil important food sources of tiny, freshwater species: *fresh detritus have protrusible pharynx or a nonprotrusible pharynx with or without jaw pharynx is highly muscular, line internally and externally with cuticle and has a triangular lumen mouth is situated in the anterior sucker jaw- bears small teeth along the edge; large, oval and bladelike hirudin- anticoagulant secreted by the salivary glands stomach may be a straight tube short esophagus either opens to a stomach or expands to a crop 1 to 11 pairs lateral cecca carnivores or blood sucking ecto parasites

differentiated into: *pharynx- protrusible, tongue-like, muscular bulb or an eversible organ *short esophagus *stomach *intestine *rectum bulb- situated in midventral wall of foregut teeth of various form and function occur in the pharynx ceca and anterior end of the intestine secrete digestive enzymes deposit feeders that used organic material in sediment as food for burrowers and sedentary burrow- and tube dwellers direct deposit feedersingest sediment directly with mouth

*ectodermal foregut *endodermal midgut (stomach, intestine) *ectodermal hindgut or rectum forehut is specialized into 2 regions: *muscular, often protrusible or eversible pharynx *a ciliated esophagus that links the pharynx to the midgut

*algae *other microorganism digestive tract straight and relatively simple mouth situated beneath the prostomium principal ingestive organ is a mid-dorsal pharyngeal bulb pharyngeal bulb is everted and collects particle on its adhesive surface pharynx acts as a pump pharyngeal gland produces salivary secretions containing mucus and enzymes gizzard- used for grinding food particles, lined with chitinous cuticle, very muscular

indirect deposit feedersfeed on organic material on sediments but collect the material first w/ a specialized appendage

crop- storage chamber; thin walled

carnivore, herbivore and scavengers for motile

calciferous glands- secrete calcium carbonate into the esophagus in the form of calcite crystals; do not play a role in digestion ciliated intestine- forms the remainder of the digestive tract anterior half of intestinesite of enzyme secretion and digestion; posterior half- primarily absorptive

suspension feeders for sedentary animals have a well developed muscular, eversible pharynx sometimes has thick, rasping lower lip/ it may bear jaws with grasping or venomous teeth few species are parasites readily generate missing or damaged body parts nervous system plays an important role in regeneration common modes of asexual reproduction: *budding *transverse division or fragmentation Sexual reproduction:

Reproduction and Development

primitively gonochoric spawn gamets through metanephridia clonal (asexual) reproduction: *fragmentation *budding *a form of transverse division zygotes undergo spiral cleavage and develop into trochopore larva larval episphere- becomes the peristomium

clonal reproduction is common transverse division of parent into two or more new individual some are hermaphroditic undergoes copulation direct deveopers have relatively yolky eggs

do not reproduce asexually cant regenerate lost parts hermaphrodites (protandric/not simultaneous hermaphrodites) testes mature before the ovary sperm transmission is similar to earthworm/copulation cryptolarva develop within the coccoon have annual or two year cycle, breeding in spring and maturing by the following year

majority are gonochoric gametes are spawned by metanephridia or by body wall rupture

larva is absent development into juvenile occurs in the cocoon

body posterior to the telotorch- becomes the pygidium

Epitoky- formation of a pelagic reproductive individual/epitoke from a benthic non-reproductive individual/atoke swarming- many epitoke swim to the surface and shed eggs and sperm female produces pheromone that attracts male and stimulate shedding of sperm sperm stimulate the shedding of egg swarming is affected by: *light cues *lunar periods embryo develops into a top shaped trocophore larva after gastrulation trocophores are lecitotrophic metamorphosis transforms trocophore to the juvenile body