Chapter 12 SMALLER ECDYSOZOANS 12-1 12-2 Diversity • Many protostomes possess a cuticle – Non-living outer layer secreted by epidermis • Cuticle restricts growth and must be molted via ecdysis • Members of Ecdysozoa molt cuticle as they grow • Regulation of molting achieved by the hormone ecdysone 12-3 12-4 Phylum Nematoda: Roundworms • 12,000 species have been named – As many as half a million may exist • Found in virtually all habitats in all biomes – Topsoil may contain billions per acre • Nematode parasites exist in nearly all animal and plant species 12-5 Phylum Nematoda: Roundworms Form and Function Outer body covering is a thick, noncellular cuticle, secreted by the underlying epidermis- hypodermis Cuticle serves to contain the hydrostatic pressure exerted by fluid in the pseudocoelom • Collagen is the primary protein in layers of the cuticle • Muscles – Longitudinal muscles lie beneath the cuticle • No circular muscles – Run in four bands, marked off by epidermal cords that project inward to pseudocoelom – Unlike other animals, the muscle extends to the nerve cords 12-6 12-7 Phylum Nematoda: Roundworms • Hydrostatic skeleton-muscle contractions press on fluid • No circular muscles to compliment longitudinal muscle in movement (antagonist) so the cuticle assists. • Fluid is force to opposite side when longitudinal muscles constrict forcing the cuticle to expand • Compression and stretching of the cuticle returns the body to resting position when muscles relax • Produces the thrashing movement of nematodes 12-8 Phylum Nematoda: Roundworms • Digestion – Gut tube consists of mouth, pharynx, non-muscular intestine, short rectum and anus – Muscular pharynx sucks food in – Intestinal wall is one cell thick-no muscles • Food moves back as new food enters and the body moves – Defecation occurs from opening the anus and allowing pseudocoelomic pressure to expel waste • Some parasitic adults have an anaerobic metabolism; aerobic metabolism are absent • Free-living nematodes and free-living stages of parasitic nematodes have both anaerobic and aerobic metabolism 12-9 Phylum Nematoda: Roundworms • Ring of nerve tissue and ganglia around the pharynx give rise to small nerves to the anterior end and to two nerve cords, one dorsal and one ventral • Sensory organs at head and tail • Most are dioecious with males smaller than females • Male has copulatory spicules • Fertilization is internal • Nematode sperm has no flagella, in female reproductive tract sperm is ameboid and moves by pseudopods • Eggs are stored in uterus until deposited • Cuticle is shed between each of four juvenile stages 12-10 Figure 12_03 Representative Nematode Parasites Representative Nematode Parasites • Some are parasites of humans – Most are tropical • Ascaris lumbricoides – Occurs in up to 25% of people in some areas of the southeastern U.S. – More than 1.27 billion affected worldwide – A. suum is found in pig intestines – A female Ascaris may lay 200,000 eggs a day, which pass out in host’s feces 12-12 Figure 12_04a Representative Nematode Parasites – Survive for long periods in soil – When humans eat undercooked vegetables contaminated with shelled juveniles, or when children put soiled fingers or toys in their mouths, consumed juveniles hatch, and burrow through intestinal wall – Carried through the heart to the lungs, they enter into alveoli causing pneumonia and are carried up to tracheae – Coughed up and swallowed, they mature in the intestine – They feed on intestinal contents and may block the intestines 12-14 Figure 12_04b Representative Nematode Parasites • Hookworms – – – – Anterior end of these small worms has a hook-like curve Necator americanus, most common hookworm. Sexes are separate Large plates in mouth cut into intestinal mucosa and suck host’s blood – Pump through more blood than they digest • Heavy infections cause anemia – Eggs pass out in feces and juveniles hatch in soil – If human skin comes in contact with soil, infective juveniles burrow through skin to blood – Travel in blood to the lungs, are coughed up to be swallowed, and mature in the intestine 12-16 Figure 12_05 Representative Nematode Parasites • Trichina Worm – Trichinella spiralis causes a potentially lethal trichinosis – Adult worms burrow into intestinal mucosa of the host’s small intestines and females directly produce juvenile worms – Juveniles penetrate blood vessels and circulate throughout the body to all tissues and spaces – Penetrate skeletal muscle cells, redirecting gene expression of the musculature • Cells lose striations and becomes a nurse cells to the parasite – When poorly cooked meat (pork or bear) containing encysted juveniles is swallowed, worms are liberated and mature in the intestine 12-18 Representative Nematode Parasites – Infect humans, pigs, rats, cats and dogs • Pigs can become infected eating uncooked scraps of infected meat or rats – Four other sibling species with variable distribution, freezing resistance, etc. – Heavy infections cause death – 25 cases of trichinosis are reported per year in the US 12-19 Figure 12_06 Representative Nematode Parasites • Pinworms – Most common worm parasite in the U.S. – Adults live in large intestine and cecum – Females, about 12 mm in length, migrate to anal region at night and lay eggs, causing itching – Scratching the anal region contaminates hands and bedclothes – Eggs develop rapidly and become infective within six hours at body temperature – When swallowed, hatch in duodenum and mature in large intestine – Members of this order have haploid males from unfertilized eggs – Females are diploid and come from fertilized eggs (haplodiploidy) 12-21 12-22 Representative Nematode Parasites • Filarial Worms – Eight species of filarial nematodes infect humans • Some cause serious diseases – Wucheria bancrofti and Brugia malayi live in lymphatic system • Cause inflammation and obstruction of the lymphatics vessels • Females release live young, tiny microfilariae, into blood and lymph • Mosquitoes ingest microfilariae when they feed • Worms develop to infective stage and move into the mosquito bite wound when it bites a human • Elephantiasis is caused by repeated exposure – Swelling and growth of connective tissue causes enormous swelling of body parts-scrotum, legs, arms 12-23 Representative Nematode Parasites – River blindness or onchocerciasis is carried by black flies and infects 37 million people in tropics – Dog heartworm, Dirofilaria immitis, is carried by mosquitoes and is the most common U.S. filarial worm – Heartworm pills 12-24 Figure 12_08 Figure 12_09 Representative Nematode Parasites – Dracunculus medinensis, the guinea worm - Larvae live within planktonic copepods (water fleas) - Upon ingestion of contaminated water, water fleas are digested by human host, but larvae survive and penetrate the stomach/intestinal wall - Worms live and mate within body cavity - Gravid females migrate through tissues to the lower extremities and produce an open ulcer - Causing burning pain, the human host immerses leg into water and female releases thousands of eggs into water - Eggs are eaten by copepods and the cycle continues 12-27 Representative Nematode Parasites • Only known treatment is to carefully remove ulcerated females carefully with a stick; treated this way for thousands of years Phylum Nematomorpha Diversity • “Horsehair worms” resemble coarse hairs • Adult structures resemble those seen in nematodes: cuticle, epidermal cords, only longitudinal muscles, and a similar nervous system pattern – Currently placed as the sister taxon to nematodes • About 320 species are known – Occur worldwide • Adults are free-living in moist habitats • Juveniles are parasites of arthropods 12-30 Phylum Nematomorpha – Larvae encyst within host and do not emerge from an aethropod host unless water is nearby – Juveniles of freshwater forms use terrestrial insects as hosts – Marine nematomorphs infect certain crabs • Digestive system is vestigial – Larvae absorb food from arthropod hosts Adults can absorb organic molecules through vestigial gut and body wall 12-31 Figure 12_10 Phylum Tardigrada • Known as “water bears” • Very small, less than 1 mm long • About 900 live in a water film around mosses and lichens • Some live in freshwater and some are marine • Most are terrestrial that occupy a film of water surrounding mosses and lichens • Trunk bears eight short unjointed legs, each with claws • A pair of stylets and sucking pharynx protrude to pierce nematodes or plant cells • Body covered by non-chitinous cuticle that is molted four or more times during lifetime • Most of the body cavity a hemocoel • No circulatory or respiratory systems 12-33 • Cryptobiosis – Terrestrial tardigrades can suspend metabolism to survive harsh conditions – Tardigrades can dehydrate from 85% water to only 3% water • In this state they can resist extreme temperatures, ionizing radiation, oxygen deficiency, etc. for years – When water is available, they become metabolically active again 12-34 • Reproduction – Sexes are separate – In parthenogenetic freshwater and moss-dwelling species, males are unknown 12-35 Figure 12_15 Phylogeny and Adaptive Diversification Phylogeny • Evolutionary relationships among ecdysozoans are not well-understood • Members of this clade do not share a common cleavage pattern – Nematodes and nematomorphs • Cleavage is unique, not spiral or radial – Cleavage in tardigrades • Has yet to be studied 12-37 Phylogeny and Adaptive Diversification • Recent phylogenies place Nematoda and Nematomorpha as sister taxa since they share a collagenous cuticle 12-38 Phylogeny and Adaptive Diversification • Tardigrades have some similarities to rotifers, particularly in their reproduction and cryptobiotic tendencies • Tardigrades and arthropods also share arthropodtype setae and muscles inserted on the cuticle 12-39 Phylogeny and Adaptive Diversification Adaptive Diversification • • • • • Nematodes show the most impressive adaptation Found in almost every habitat available to animals Body structure is plastic enough to allow adaptation Life cycle ranges from simple to complex Have been known to survive in suboptimal conditions 12-40