NEMATODE EXCRETORY SYSTEM Section through esophagus Section through middle Stylet Median Bulb Lips Procorpus Excretory Duct Bursa Spicules in cloaca Intestine Lobe of esophagus gland overlapping intestine Testis Ovary Intestine Vulva Egg Postuterine Sac (Present in species with 1 ovary) Lateral Lines Anus Tail Basic Nematode Body Design C.I.H. 1:21 D. destructor NEMATODE EXCRETORY SYSTEM Excretory canals run through lateral hypodermal cords. Different systems = “H” (A, B), “U” (C), and tylenchoid (D). A single “renette cell” (E) is present in Dorylaimida. NEMATODE PHYSIOLOGY EXCRETORY SYSTEM Excretion of Nitrogenous Waste Products -Ammonia, urea, or uric acid -Amino acids, amines, peptides & fatty acids -Aquatic organisms = live in soil water -Ammonia toxic but diffuses away in soil water -Urea, nucleic acids, etc. Diffuse through body wall Voided through intestine, excretory system Stored in crystalline state NEMATODE PHYSIOLOGY EXCRETORY SYSTEM Other Functions -Osmoregulation -Removing excess water -Regulating turgor in body cavity -General “cleanup”, removes foreign particles -Releases enzyme-containing secretions a) Exodigestive enzymes b) Protective coatings c) Assist molting No Circulatory System No Respiratory System Circulatory and respiratory processes are accomplished by movement of fluids within the body cavity. NEMATODE PHYSIOLOGY RESPIRATION Nematodes are aerobic organisms -Require oxygen for metabolism -Thin shape and semipermeable cuticle -Width of nematode restricted by diffusion -Oxygen availability and demand depend on: 1) O2 concentration in environment 2) Diffusion of O2 into microsite 3) Metabolic demand from body size, activity, temperature, starvation NEMATODE PHYSIOLOGY RESPIRATION When oxygen is insufficient, nematodes -Decrease rate of development -Stop moving -Enter cryptobiosis NEMATODE PHYSIOLOGY RESPIRATION Many nematodes are facultative anaerobes -Persist through microaerobic (<5% O2) and anaerobic (no O2) periods with fermentative metabolism of glycogen Caenorhabditis survived 0% O2 for 80 hours Aphelenchus survived >30 days! NEMATODE PHYSIOLOGY RESPIRATION Oxidative metabolism preferred 1) More energy produced 2) Toxic waste products produced from fermentative metabolism O2 uptake is reduced as O2 levels decline from normal air (21%) to 5%. At <5% O2, O2 uptake stops and nematodes must switch to fermentative metabolism NEMATODE PHYSIOLOGY MOLTING As nematodes grow they need to shed their cuticle to increase in size. Nematodes generally go through 4 molts. J1 J2 J3 J4 Adult Timing of molting depends on “stage of development”(physiological age), not necessarily chronological age, so each stage tends to be a certain size for a given species. NEMATODE PHYSIOLOGY MOLTING Just before molting begins, the nematode stops feeding and becomes sluggish or inactive. A = Normal cuticle. B = Initial stages of molting, the hypodermis thickens C = Hypodermis secretes new cuticle under the old one NEMATODE PHYSIOLOGY MOLTING D = Two cuticles separate and a space develops between them. Folds may appear in the old cuticle as it enlarges and becomes loose. E = Most of the old cuticle is reabsorbed before it is shed. F = The old external cortex is not reabsorbed and is the only part of the cuticle cast off. NEMATODE PHYSIOLOGY MOLTING The nematode breaks out of the old cuticle by "rubbing" against soil particles. New cuticle is convoluted to allow for rapid growth, increase in thickness. The entire cuticle is shed including the lining of the stoma, esophagus, vulva, cloaca, & rectum. New cuticular structures are formed. The basal part of the old stylet dissolves and slow retractile movements disengage the head from the anterior part of the old stylet, which remains attached to the anterior part of the old cuticle. R.S. Hussey NEMATODE PHYSIOLOGY SENESCENCE, QUIESCENCE & CRYPTOBIOSIS Normal life spans for most individuals of a given species are similar. Normal aging process: = Senescence NEMATODE PHYSIOLOGY SENESCENCE, QUIESCENCE & CRYPTOBIOSIS When environment shifts unfavorably nematodes experience metabolic “slowdown” = Quiescence Movement slows or stops, life cycle development delayed. Metabolism slowed, but still detectable. NEMATODE PHYSIOLOGY SENESCENCE, QUIESCENCE & CRYPTOBIOSIS Increased or prolonged environmental stress leads to metabolic “shutdown” = Cryptobiosis Metabolism is not detectable! NEMATODE PHYSIOLOGY SENESCENCE, QUIESCENCE & CRYPTOBIOSIS Physiological aging slows in quiescent nematodes. 64-day-old quiescent Meloidogyne javanica = physiological age of 4 days. 128-day-old Tylenchulus semipenetrans had aged 16 days. Aphelenchus avenae under O2 stress for 90 days had not aged at all. NEMATODE PHYSIOLOGY SENESCENCE, QUIESCENCE & CRYPTOBIOSIS Several environmental conditions may induce nematodes to enter a quiescent or cryptobiotic state. Anhydrobiosis = condition due to lack of water. Cryobiosis = caused by lowering of temperature. Anoxybiosis = absence or low levels of oxygen. Osmobiosis = removal of water from organisms by a solution that has high osmotic potential. Several may occur a same time. Anhydrobiosis most common. NEMATODE PHYSIOLOGY SENESCENCE, QUIESCENCE & CRYPTOBIOSIS Induction of anhydrobiosis due to slow dehydration includes physical and physiological responses to resist desiccation. Physical response = coiling into a spiral reduces the surface area exposed to the environment and reduces the rate of water loss. When dried straight, nematodes have little control of water loss, lose water too quickly. Aphelenchus avenae in anhyrobiosis Coiling = physical response M.A. McClure NEMATODE PHYSIOLOGY SENESCENCE, QUIESCENCE & CRYPTOBIOSIS Some nematodes swarm into a mass that also helps protect them from rapid water loss. At times, a lipid layer surrounding the cuticle causes the swarm to clump and stick together (= nematode wool). Nematodes inside plant tissues are further protected. Swarm of Anquina tritici juveniles in anhydrobiotic state M.A. McClure Ditylenchus dipsaci J4 – nematode wool on basal plate defra defra NEMATODE PHYSIOLOGY SENESCENCE, QUIESCENCE & CRYPTOBIOSIS Physiological Responses – 1. Water Dynamics Active nematode = 75-80% water. Anhydrobiotic in dry air (0% rh) is 2-5% water. Water lost by dehydration (80% -> 5%) is "free water" = used in metabolic processes and in cytoplasm. Can be lost without cell damage. The 2-5% water remaining is "bound water" and is essential for the maintenance of membranes and other structural components. NEMATODE PHYSIOLOGY SENESCENCE, QUIESCENCE & CRYPTOBIOSIS Physiological Responses – 2. Metabolic Changes During dehydration - lipid concentration decreases - trehalose (a non-reducing sugar) increases 7X - glycerol concentration increases substantially Glycerol replaces bound water in stabilizing membrane structure. "Quick dried" nematodes do not synthesize glycerol, membranes destabilize, and nematode dies. NEMATODE PHYSIOLOGY SENESCENCE, QUIESCENCE & CRYPTOBIOSIS Resistance to Unfavorable Conditions Cryptobiotic = resistant to harsh environment Don’t want conditions that would induce cryptobiosis before you treat! Examples: See Handout NEMATODE PHYSIOLOGY RESISTANT STAGES In some nematodes only one life stage from egg to adult is resistant. Can be any stage. See handout for examples In other species, any stage from the J2 to adult can successfully enter and exit cryptobiosis. NEMATODE PHYSIOLOGY RESISTANT STAGES Another form of resistance strategy is called the dauer juvenile (= dauer larvae). The third stage juvenile retains the second stage cuticle = double cuticle. Common among bacterial feeders. The body is usually thinner, cuticle thicker, and nematode does not feed. This stage is often used during migration. NEMATODE PHYSIOLOGY NEMATODE LONGEVITY Nematodes can persist in a resistant state for many years. Normal life spans of active nematodes Caenorhabditis = 4 days Xiphinema = 3 years Most plant parasites ≈ 35 days Longer if cooler, shorter if warmer. Longevity is more an expression of physiological state than of time.
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