Innate immunity of fish (overview).

The innate immune system of fish is considered to be the first line of defence against a broad spectrum of pathogens and is more important for fish as compared with mammals. Lysozyme level or activity is an important index of innate immunity of fish and is ubiquitous in its distribution among living organisms. It is well documented that fish lysozyme possess lytic activity against both Gram-positive bacteria and Gram-negative bacteria. It is also known to be opsonic in nature and activates the complement system and phagocytes. It is present in mucus, lymphoid tissue, plasma and other body fluids of freshwater and marine fish. It is also expressed in a wide variety of tissues. Lysozyme activity has been shown to vary depending on the sex, age and size, season, water temperature, pH, toxicants, infections and degree of stressors. Here, we review our current understanding of different types of lysozyme and their expression and its role in fish innate immune system.

Lysozyme: an important defence molecule of fish innate immune system

Stress is an event that most animals experience and that induces a number of responses involving all three regulatory systems, neural, endocrine and immune. When the stressor is acute and short-term, the response pattern is stimulatory and the fish immune response shows an activating phase that specially enhances innate responses. If the stressor is chronic the immune response shows suppressive effects and therefore the chances of an infection may be enhanced. In addition, coping with the stressor imposes an allostatic cost that may interfere with the needs of the immune response. In this paper the mechanisms behind these immunoregulatory changes are reviewed and the role of the main neuroendocrine mechanisms directly affecting the building of the immune response and their consequences are considered.

Stress and immune modulation in fish.

All metazoans possess innate immune defence system whereas parameters of the adaptive immune system make their first appearance in the gnathostomata, the jawed vertebrates. Fish are therefore the first animal phyla to possess both an innate and adaptive immune system making them very interesting as regards developmental studies of the immune system. The massive increase in aquaculture in recent decades has also put greater emphasis on studies of the fish immune system and defence against diseases commonly associated with intensive fish rearing. Some of the main components of the innate and adaptive immune system of fish are described. The innate parameters are at the forefront of immune defence in fish and are a crucial factor in disease resistance. The adaptive response of fish is commonly delayed but is essential for lasting immunity and a key factor in successful vaccination. Some of the inherent and external factors that can manipulate the immune system of fish are discussed, the main fish diseases are listed and the pathogenicity and host defence discussed. The main prophylactic measures are covered, including vaccination, probiotics and immunostimulation. A key element in the immunological control of fish diseases is the great variation in disease susceptibility and immune defence of different fish species, a reflection of the extended time the present day teleosts have been separated in evolution. Future research will probably make use of molecular and proteomic tools both to study important elements in immune defence and prophylactic measures and to assist with breeding programmes for disease resistance.

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Immunological Control of Fish Diseases

Tissue trauma or invasion by pathogens or parasites induce changes in the quantities of several macromolecules in animal body fluids These changes comprise one aspect of the acute phase response (APR), which in toto involves metabolic changes in several organ systems One clear indication of the response is the increase in synthesis and secretion by the liver of several plasma proteins, with simultaneous decreases in others These acute phase proteins (APP) function in a variety of defense-related activities such as limiting the dispersal of infectious agents, repair of tissue damage, inactivation of proteases, killing of microbes and other potential pathogens, and restoration of the healthy state Some APP are directly harmful to microbes, while others modify targets thus marking them for cell responses Some work alone while others contribute to cascades Proteins that are APP in mammals, and that have been identified in both teleosts and elasmobranchs include C-reactive protein, serum amyloid P, and several components of the Complement system Others reported in teleosts include transferrin and thrombin Of these, only CRP has been reported to increase in acute phase plasma In trout, a precerebellin-like protein is an APP with unknown functions A cDNA library enriched in fragments of transcripts that were more abundant in livers from fish undergoing an APR recently yielded sequences resembling 12 additional known APP, and as many others either not known to be APP, or not similar to others yet in public databases It appears that, as in mammals, hepatocytes are the prime source of APP in fish, and that pro-inflammatory cytokines induce transcription of their genes

The acute phase response and innate immunity of fish

The fight-or-flight response prepares an animal for coping with alarming situations and their potential consequences, which include injury. The possible involvement of innate components of immunity in the response has received little attention. We determined plasma concentrations of stress hormones and lysozyme activity before and after a 10 min handling stressor. Rainbow trout (Oncorhynchus mykiss) were anesthetized in their home tanks, bled, revived, and then stressed by being held in the air in a net for 30 s and placed in a shallow bucket of water for 10 min. Fish were then captured, concussed (in one of two experiments) and bled again. Control fish were also bled twice, but were kept anesthetized in their holding tanks between bleedings. Following the stressor, plasma cortisol, adrenaline and lysozyme activity were significantly increased. The experiment was repeated 4 months later with a similar outcome. While chronic stress is eventually immunosuppressive, acute stress/trauma may help enhance both cellular and humoral components of innate defenses at times of likely need.

The immediate effects of stress on hormones and plasma lysozyme in rainbow trout

Sympathetic activation leading to increased levels of blood catecholamines, and stimulation of the hypothalamic‐pituitary‐inter-renal axis leading to increased cortisol, are difficult to avoid when handling animals. Yet, in research on effects of acute stress, elicitation of such responses must be minimized in the control groups. The work examines means to achieve a minimally disturbed state in rainbow trout (Oncorhynchus mykiss). Level of arousal was determined by adrenaline and cortisol concentrations in plasma, and by the spleen:somatic index. Fish were prepared for bleeding by rapid capture and concussion, by infusion of anesthetic into the undisturbed home tank, by confinement in black boxes, or by being fed a- and b-receptor antagonists. Even when done quickly, netting and concussion yielded fish with ca. 200-pmol adrenaline:ml plasma. Cortisol was elevated (to \10 ng:ml) within 30 s of stress initiation. Surreptitious infusion of anesthetic (2-phenoxyethanol, PE) into tanks yielded fish with lower adrenaline levels (means 19.34 and 19.58 pmols:ml in home tank and black boxes, respectively). Among fish given phentolamine and propranolol, spleen:somatic indices and plasma adrenaline were higher than in diet controls, whether undisturbed or stressed, indicative of successful receptor blockade. Since careful infusion of 2-PE yielded the lowest adrenaline levels, and requires no special apparatus, it is the method of choice for obtaining minimally stressed fish. © 1999 Elsevier Science Inc. All rights reserved.

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Nora E. Demers

Papers

The immediate effects of stress on hormones and plasma lysozyme in rainbow trout

Modulation of stress hormones in rainbow trout by means of anesthesia, sensory deprivation and receptor blockade