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Fish gill

Fish gills are the primary respiratory organs in most fish species, consisting of specialized, feathery structures located in the pharyngeal cavity that enable the extraction of dissolved oxygen from water and the expulsion of carbon dioxide into it through diffusion across a thin epithelial barrier.^[1] These organs are typically protected by a bony operculum and comprise four pairs of gill arches in bony fishes,^[2] each supporting numerous gill filaments that branch into secondary lamellae to maximize surface area for gas exchange, often reaching 0.1–0.4 m² per kilogram of body weight.^[1] Beyond respiration, fish gills serve multifunctional roles, including osmoregulation through active ion transport via mitochondrion-rich cells (such as chloride cells) to maintain salt and water balance in varying salinities, acid-base regulation by secreting protons or bicarbonate ions, and the excretion of nitrogenous wastes like ammonia, which accounts for over 80% of such elimination in many species.^[1] The efficiency of these processes is enhanced by a countercurrent exchange mechanism, where blood flows opposite to water across the gill lamellae, allowing oxygen uptake rates of 50–90% from ambient water.^[1] Gill rakers, comb-like projections on the arches, additionally aid in filter-feeding by trapping food particles while preventing debris from damaging the delicate filaments.^[3]

Anatomy and Structure

Gill Arches

Gill arches form the foundational skeletal framework of fish gills, consisting of curved structures composed of bone in bony fishes or cartilage in cartilaginous fishes, typically arranged in four pairs (eight arches total, with four on each side of the head).^[4]^[5] These arches are positioned posterior to the mouth and buccal cavity, within the pharyngeal region, where their ends attach to the interbranchial septa that divide the gill pouches.^[3]^[6] The primary role of gill arches is to suspend the delicate gill filaments, providing structural support that maintains the overall architecture of the gill for effective function.^[7] Additionally, these arches serve as key attachment points for muscles, such as the dorsal gill arch muscles, which facilitate subtle movements of the gills during ventilation by anchoring the arches to the cranium and other skeletal elements.^[8] This muscular attachment helps stabilize the gill apparatus against water flow pressures.^[9] While four pairs predominate in most teleost fishes, variations occur across species; for instance, some primitive bony fishes exhibit five pairs, cartilaginous fishes typically have 5–7 pairs, and jawless fishes possess seven pairs (fourteen arches total, with seven on each side).^[10] Rigidity also differs, with bony arches being more robust in advanced teleosts compared to the flexible cartilaginous forms in sharks and rays, adaptations reflecting diverse habitats and feeding strategies.^[4]

Gill Filaments and Lamellae

Gill filaments, also known as primary lamellae, are elongated, finger-like projections that extend from the gill arches, serving as the primary sites for gas exchange in fish gills. These structures are typically arranged in rows along the arches, with each filament measuring several millimeters in length and containing numerous secondary lamellae. Secondary lamellae, the finer subdivisions, appear as thin, plate-like folds on the surface of the primary lamellae, greatly increasing the gill's total surface area available for diffusion. The lamellae are composed primarily of a thin epithelial layer that facilitates the diffusion of gases and ions, supported internally by specialized cells. Epithelial pavement cells form the outer covering, providing a barrier that is only a few micrometers thick to minimize diffusion distance. Pillar cells, unique to fish gills, create a lattice-like framework within the lamellae, forming channels that maintain structural integrity and support blood flow. Chloride cells, or ionocytes, are interspersed among the epithelial cells, particularly on the afferent side, and are responsible for ion regulation and osmoregulation. In active fish species, such as salmonids, the gill surface area can reach estimates of 0.1 to 0.2 m² per kg of body weight, enabling efficient oxygen uptake during high metabolic demands. This expansive area is crucial for aquatic respiration, where oxygen availability is lower than in air. To prevent collapse under water flow or pressure changes, interlamellar water channels exist between secondary lamellae, providing mechanical support and allowing continuous water passage. These filaments attach to the gill arches to ensure overall structural stability during ventilation.

Blood Supply and Countercurrent Exchange

Deoxygenated blood from the heart reaches the gills via the ventral aorta, which branches into four pairs of afferent branchial arteries, one for each gill arch. These arteries supply deoxygenated blood to the afferent filament arteries running along the leading edge of each gill filament. Within the secondary lamellae, blood enters narrow channels via afferent lamellar arterioles at the base of the lamella, flows through the capillary network for gas exchange, and exits via efferent lamellar arterioles to the efferent filament arteries along the trailing edge of the filament. Oxygenated blood then converges into the efferent branchial arteries, which unite to form the dorsal aorta for systemic distribution. The efficiency of gas exchange in fish gills relies on a countercurrent flow arrangement, where blood and water move in opposite directions across the lamellar surface.^[11] Water enters the gill slits and flows over the lamellae from the afferent (leading) edge toward the efferent (trailing) edge, progressively losing oxygen.^[11] Simultaneously, deoxygenated blood enters the lamellae at the afferent side, where it encounters relatively oxygen-rich water, and flows toward the efferent side, where it meets oxygen-depleted but still higher-oxygen water compared to the blood itself.^[11] This opposing flow maintains a consistent concentration gradient along the entire length of the lamella, enabling up to 80-90% oxygen extraction from water, far exceeding the 50% limit of concurrent flow systems.^[11] In a diagrammatic cross-section of a lamella, arrows would depict water flowing left to right over the surface while blood capillaries run right to left beneath, ensuring the blood's oxygen content rises progressively as water's falls. The rate of oxygen diffusion across the gill epithelium follows Fick's law of diffusion, expressed as:

J = -D \cdot \frac{\Delta C}{\Delta x}

where J is the diffusion flux, D is the diffusion coefficient of oxygen in the tissue barrier, \Delta C is the concentration difference across the barrier, and \Delta x is the barrier thickness.^[12] In fish gills, the countercurrent system sustains a favorable \Delta C by preventing equilibrium, while thin barriers (typically 0.5-1 μm) and high surface area minimize \Delta x and maximize flux; adaptations like reduced cell size further optimize D.^[12] Pillar cells, specialized flattened endothelial cells unique to fish gills, form the structural framework of the lamellar blood channels.^[13] These cells extend flange-like processes that create interdigitating blood lacunae separated by collagen bundles, ensuring a thin, uniform diffusion distance while supporting high blood pressure without collapse.^[13] By containing contractile actomyosin filaments, pillar cells regulate channel diameter, preventing blood swelling or pooling that could increase diffusion path length and impair exchange efficiency.^[14]

Respiratory Function

Ventilation Mechanism

Fish primarily ventilate their gills through a buccal-opercular pumping mechanism, which involves coordinated expansion and contraction of the mouth (buccal cavity) and opercular chamber to generate pressure gradients that drive water flow.^[15] During the inspiratory phase, the mouth opens while the opercula remain closed, and the buccal cavity expands slightly ahead of the opercular cavity, creating negative pressure that draws water into the mouth.^[3] In the expiratory phase, the mouth closes, the opercula open, and both cavities contract, with the buccal contraction leading to produce positive pressure that forces water unidirectionally over the gill surfaces and out through the opercular slits.^[15] This pumping action ensures unidirectional water flow from the mouth through the gill arches to the opercular chamber, preventing backflow and mixing of incoming fresh water with outgoing deoxygenated water, which maintains efficient oxygen extraction.^[3] The process allows water to pass over the gills in a consistent direction, facilitating the gas exchange that occurs as oxygen diffuses into the blood.^[15] The energy cost of this active ventilation is substantial, accounting for approximately 10-15% of the total oxygen uptake in resting or slowly swimming fish, reflecting the metabolic investment required to overcome gill resistance and sustain pumping.^[16] In fast-swimming species such as lamnid sharks and tunas, an adaptation known as ram ventilation replaces or supplements pumping; these fish swim continuously with mouths agape, using forward momentum to ram water over the gills without muscular effort from the buccal or opercular regions.^[17] This obligate ram ventilation is efficient at high speeds but requires constant motion to prevent suffocation.^[17]

Gas Exchange Process

The gas exchange process in fish gills primarily involves the passive diffusion of oxygen (O₂) and carbon dioxide (CO₂) across the thin lamellar epithelium, driven by partial pressure gradients between the surrounding water and the blood. Oxygen diffuses from water into the blood when the partial pressure of O₂ (PO₂) in the ventilated water exceeds that in the deoxygenated blood entering the gills, while CO₂ diffuses in the opposite direction due to its higher partial pressure (PCO₂) in the blood compared to the water.^[18] This diffusion adheres to Fick's law, where the rate is proportional to the surface area, diffusion coefficient, and partial pressure difference, but is fundamentally influenced by Henry's law, which states that the solubility of a gas in a liquid is directly proportional to the partial pressure of that gas above the liquid.^[19] In the context of gills, Henry's law explains why O₂ solubility in water is low (approximately 7 ml/L at standard conditions, about 3% of its solubility in air), creating a steeper diffusion gradient compared to blood, where hemoglobin enhances O₂ carrying capacity; conversely, CO₂ has higher solubility in water than in blood, facilitating its excretion despite lower overall diffusion rates for CO₂ due to bicarbonate transport limitations.^[18] The efficiency of gas exchange is heavily influenced by the gill's surface area-to-volume ratio, which determines the available interface for diffusion; higher ratios in active species enhance oxygen uptake rates to support elevated metabolic demands. For instance, rainbow trout (Oncorhynchus mykiss), a highly active fish, possess a respiratory gill surface area of approximately 550 cm² per 300 g body weight, with the vascular area nearly 1.65 times larger—allowing efficient O₂ extraction even during sustained swimming.^[18] In contrast, sluggish species like carp exhibit lower surface area-to-volume ratios, resulting in reduced exchange rates and adaptations for lower activity levels.^[20] Environmental factors such as pH and temperature modulate diffusion rates in fish gills by altering gas solubility and metabolic kinetics. According to Henry's law, rising temperatures decrease O₂ solubility in water (e.g., by 20-30% per 10°C increase), steepening partial pressure gradients but potentially limiting overall availability unless compensated by increased ventilation; in rainbow trout, acute warming to 20°C from 10°C induces gill remodeling to boost surface area and sustain diffusion.^[18] pH shifts affect CO₂ diffusion indirectly by influencing bicarbonate equilibria and acid-base regulation, with acidic conditions (pH <7) accelerating HCO₃⁻/Cl⁻ exchange to enhance CO₂ unloading, while alkaline pH reduces ammonia diffusion across the epithelium.^[18] Countercurrent flow between water and blood continuously maintains these gradients for maximal efficiency.^[18]

Oxygen Affinity and Regulation

Fish hemoglobin exhibits a higher oxygen affinity compared to human hemoglobin, facilitating efficient oxygen loading from the low partial pressures typical in aquatic environments. The P50 value, representing the partial pressure of oxygen at which hemoglobin is 50% saturated, for fish blood generally ranges from 20 to 40 mmHg, lower than the approximately 27 mmHg observed in humans at standard conditions of pH 7.4 and 37°C.^[21]^[22]^[23] This elevated affinity ensures near-complete saturation of hemoglobin in the gills despite ambient oxygen levels often below 100 mmHg.^[22] The Bohr effect plays a crucial role in regulating oxygen affinity within fish gills by modulating the hemoglobin-oxygen dissociation curve in response to respiratory gases and pH. As carbon dioxide diffuses into the blood during gas exchange, it lowers the pH, reducing hemoglobin's oxygen affinity and promoting unloading at tissues; conversely, at the gills, deoxygenation and CO2 release facilitate higher affinity for incoming oxygen.^[24]^[25] This pH-dependent shift, quantified as Δlog P50/ΔpH (typically -0.4 to -0.6 in teleosts), optimizes oxygen transport efficiency across the respiratory cycle.^[24] In certain fish species, such as carp and trout, the Root effect represents an extreme manifestation of pH sensitivity, where hemoglobin saturation decreases even at high oxygen partial pressures under acidic conditions. This phenomenon, characterized by a pronounced reduction in maximum oxygen-carrying capacity (e.g., saturation dropping below 50% at pH 6 despite PO2 > 100 mmHg), aids in oxygenating specialized structures like the swim bladder and retina.^[26]^[22] The Root effect enhances overall oxygen delivery potential by up to 73% in species like trout, beyond the contributions of the standard Bohr effect.^[26] Hormonal regulation further fine-tunes gill oxygen uptake through modulation of blood flow, particularly via catecholamines released during stress. Adrenaline and noradrenaline, secreted from chromaffin cells, act on α-adrenoceptors to induce vasoconstriction in non-respiratory gill pathways, thereby redirecting blood flow to increase perfusion through the respiratory lamellae and enhancing oxygen transfer rates.^[27] This adrenergic response, triggered by acute stressors like hypoxia, supports elevated metabolic demands by optimizing the balance between gas exchange and osmoregulation in the gills.^[27]

Variations by Fish Group

Bony Fish Gills

Bony fish, or Osteichthyes, represent the most diverse group of vertebrates, encompassing over 30,000 species, with their gills exhibiting specialized structures that support a wide range of aquatic lifestyles from freshwater streams to oceanic depths.^[28] In teleosts, the dominant subgroup, the gills consist of four pairs of bony gill arches, each bearing holobranchs—complete sets of filaments on both the anterior and posterior sides—allowing for maximized surface area within the protected gill chamber.^[28] This configuration contrasts with partial hemibranchs in some other groups and facilitates efficient gas exchange across diverse metabolic rates.^[29] The gills of bony fish are enclosed by the operculum, a bony flap that covers the gill slits and plays a crucial role in the buccal-opercular pumping mechanism for ventilation.^[30] During inhalation, the oral cavity expands to draw water over the gills, while exhalation involves opercular abduction to expel water, creating a unidirectional flow that enhances oxygen extraction without reliance on constant swimming.^[31] This protected setup is particularly advantageous for demersal or inactive species, enabling sustained respiration in low-flow environments.^[32] In active swimmers such as salmon (Salmo salar), the gills feature high lamellar density on the secondary lamellae of filaments, providing an expansive surface area to meet elevated metabolic oxygen demands during migration and burst swimming.^[33] For instance, gill surface area in salmon scales proportionally with body mass and activity levels, supporting up to several-fold increases in oxygen uptake without compromising efficiency.^[34] This adaptation underscores the gills' role in fueling high-energy lifestyles prevalent in many teleost lineages. Freshwater bony fish exhibit distinct gill modifications, including a higher density of chloride cells (also known as ionocytes) in the lamellar epithelium to facilitate active ion uptake against osmotic gradients.^[35] These mitochondria-rich cells, abundant in species like tilapia (Tilapia zillii), employ mechanisms such as Na+/K+-ATPase and H+-ATPase to absorb essential ions like Na+ and Cl- from dilute environments, preventing ionic loss and maintaining homeostasis. Such adaptations are vital for euryhaline teleosts transitioning between habitats, where chloride cell proliferation can significantly increase in freshwater conditions.^[28]

Cartilaginous Fish Gills

Cartilaginous fish, including sharks, rays, and chimaeras, possess gill arches composed of cartilage rather than bone, supporting 5 to 7 pairs of external gill slits that open directly to the external environment without the protective covering of an operculum found in bony fish.^[36]^[7] These slits allow water to flow over the gills and exit externally, facilitating exposure to the surrounding medium while increasing vulnerability to physical damage. The gill filaments bear secondary lamellae that are typically rectangular in shape, arranged to maximize surface area for gas exchange without the need for active pumping mechanisms in many species; this configuration, supported by pillar cells, enables efficient diffusion across a thin epithelium. Ventilation in cartilaginous fish primarily relies on ram ventilation, where continuous forward swimming drives water through the mouth and over the gills, a strategy suited to their active predatory lifestyle and lower metabolic rates compared to many bony fish.^[36] This passive flow, combined with occasional active buccal pumping, supports oxygen uptake at rates sufficient for their reduced energetic demands, with water flow volumes reaching 5,000–20,000 ml·kg⁻¹·h⁻¹. The basic countercurrent exchange system in the lamellae ensures effective oxygen extraction despite the absence of an opercular pump. In addition to respiration, cartilaginous fish gills contribute to ionoregulation through mitochondrion-rich cells expressing Na⁺-K⁺-ATPase and other transporters for sodium uptake and chloride absorption, though they do not significantly excrete NaCl. This function is complemented by the rectal gland, a specialized organ unique to elasmobranchs that secretes excess salts as a hypersaline fluid, helping maintain osmotic balance in marine environments where urea retention by the gills plays a key role in matching environmental salinity. The rectal gland's activity, regulated by hormones like vasoactive intestinal polypeptide, offsets diffusive salt influx, allowing the gills to focus on gas exchange and selective ion handling.

Jawless Fish Gills

Jawless fish, represented by lampreys and hagfish, possess primitive gill arrangements consisting of internal pouches rather than the external slits and arches typical of more derived vertebrates. These structures facilitate gas exchange through a simplified system adapted to their burrowing and scavenging lifestyles.^[37] In lampreys, seven pairs of muscular gill pouches open externally through distinct gill slits on each side of the head, allowing water to flow unidirectionally over the internal gill filaments for oxygen extraction. Ventilation occurs via a velar pump—a muscular velum structure in the pharynx that draws water in through the mouth and expels it through the pouches—contrasting with the buccal pumping mechanism of jawed fish. This velar system supports both respiration and feeding, enabling lampreys to maintain attachment to hosts without interrupting gas exchange.^[38]^[39] Hagfish exhibit 5 to 16 pairs of internal gill pouches, varying by species, which receive water inhaled through a single nostril via the velum pump and exhale it separately through individual external openings. While these gills account for the majority of oxygen uptake (approximately 81%) and ammonia excretion (about 71%), the overall respiratory efficiency is lower than in jawed fish due to the pouches' isolated design and limited surface area, with cutaneous respiration supplementing up to 19% of oxygen needs in some conditions.^[40]^[41] The ammocoete larvae of lampreys feature a branchial basket—a sieving structure within the pharynx—that aids filter feeding on microorganisms while facilitating ventilation through the same velar mechanism, highlighting an integrated role for gills in early development.^[39] These pouch-like gills represent basal vertebrate traits, providing insights into the evolutionary origins of aquatic respiration and linking jawless lineages to ancestral chordates through their direct environmental exposure and lack of protective covers. Recent genomic studies, including the 2024 hagfish genome sequencing, confirm the monophyly of cyclostomes (lampreys and hagfish) and highlight conserved genetic elements potentially underlying primitive gill functions.^[42]

Additional Roles and Adaptations

Non-Respiratory Functions

Fish gills serve multiple physiological roles beyond gas exchange, leveraging their extensive surface area and specialized epithelial cells for processes such as ionoregulation, nitrogenous waste excretion, acid-base homeostasis, and sensory detection. These functions are facilitated by ion-transporting cells, including pavement cells and mitochondria-rich cells (also known as ionocytes), which express key enzymes and transporters. Ionoregulation is a primary non-respiratory function of fish gills, enabling adaptation to varying salinities through active transport of ions. In freshwater environments, where the external medium is hypoosmotic, gills actively uptake Na⁺ and Cl⁻ to counteract diffusive losses, primarily via apical Na⁺/H⁺ exchangers and basolateral Na⁺/K⁺-ATPase pumps in ionocytes. In seawater, hyperosmotic conditions drive active extrusion of Na⁺ and Cl⁻ through similar mechanisms, including Cl⁻/HCO₃⁻ exchangers and Na⁺/K⁺-ATPase, preventing ion overload. Distinct isoforms of Na⁺/K⁺-ATPase, such as α1a in freshwater and α1b in seawater, allow euryhaline species like salmon to acclimate rapidly to salinity changes.^[43]^[44] Ammonia excretion represents another critical role, as gills eliminate the primary nitrogenous waste product from protein metabolism. In most teleost fish, ammonia diffuses across the gill lamellae predominantly as un-ionized NH₃ down a favorable partial pressure gradient from blood to water, facilitated by the high permeability of the gill epithelium to this neutral form. This process accounts for over 90% of total ammonia excretion in many species, with Rh glycoproteins aiding NH₃ transport in some cases; the acidification of gill water by H⁺-ATPase traps excreted NH₃ as NH₄⁺, enhancing efficiency. In freshwater, passive NH₃ diffusion predominates, while in seawater, active components may contribute under high loads.^[45]^[46]^[47] Gills also contribute to acid-base balance by modulating plasma pH through ion exchanges and bicarbonate handling. In response to elevated blood CO₂ levels causing respiratory acidosis, gill ionocytes increase HCO₃⁻ secretion into the plasma via Cl⁻/HCO₃⁻ exchangers, while excreting H⁺ or Cl⁻ to restore equilibrium. Carbonic anhydrase enzymes in gill cells accelerate the conversion of CO₂ to HCO₃⁻ and H⁺, supporting this regulation; for instance, during hypercapnia, net HCO₃⁻ uptake or retention occurs to buffer acidosis. This gill-mediated compensation can restore blood pH within hours in species like the rainbow trout.^[48]^[49]^[50] Sensory functions of fish gills involve chemoreceptors and taste buds embedded in the gill arches and filaments, allowing detection of environmental cues. These structures house solitary chemosensory cells and taste receptor cells that respond to amino acids, ions, and pH changes, relaying signals via cranial nerves IX and X to the brain for reflexive behaviors like feeding or avoidance. In zebrafish, for example, Merkel-like cells within gill taste buds act as hypoxia chemoreceptors, triggering ventilatory adjustments through taste-signaling pathways. Such sensory capabilities enhance survival by monitoring water quality and prey.^[51]^[52]^[53]

Environmental Adaptations

Fish gills exhibit remarkable plasticity in response to hypoxic conditions, enabling enhanced oxygen uptake through morphological remodeling. In species such as crucian carp (Carassius auratus), chronic hypoxia triggers the reduction of interlamellar cell mass (ILCM), thereby exposing a greater surface area of gill lamellae for gas exchange. This adaptive remodeling increases the functional respiratory surface by up to 80% without altering the overall gill architecture, allowing the fish to maintain aerobic metabolism in oxygen-depleted waters.^[54] Similarly, some tropical fish remodel their gills by proliferating pillar cells and thinning the blood-water barrier, further optimizing diffusion efficiency under low oxygen levels.^[55] Certain fish have evolved bimodal respiration as a complementary adaptation to hypoxia, utilizing both aquatic and aerial oxygen sources. Labyrinth fish, such as the climbing perch (Anabas testudineus), possess a specialized labyrinth organ—a vascularized air-breathing structure derived from gill arches—that supplements gill ventilation during air exposure in severely hypoxic environments. This allows them to survive prolonged periods of water stagnation by partitioning respiration, with air breathing contributing up to 90% of total oxygen uptake in extreme conditions.^[56] Such adaptations are prevalent in stagnant tropical waters where dissolved oxygen can fall below 1 mg/L.^[57] In euryhaline species like the Japanese eel (Anguilla japonica), gills demonstrate plasticity to accommodate salinity shifts through targeted cell proliferation. During transfer from freshwater to seawater, chloride cells in the gill epithelium proliferate and differentiate, increasing their density by 2-3 fold to enhance NaCl secretion and maintain osmotic balance. This involves signaling pathways like MAPK and osmosensors that regulate cell turnover, enabling the gills to switch from ion uptake to excretion modes within days.^[58] In eels, this proliferation correlates with enlarged apical pits on chloride cells, facilitating active ion transport and preventing osmotic shock.^[59] Pollutants elicit defensive responses in fish gills, primarily through mucus hypersecretion and subsequent hyperplasia to mitigate toxin exposure. Heavy metals and acidic effluents induce mucous cells to release copious glycoproteins, forming a protective barrier that traps and precipitates contaminants on the gill surface, thereby reducing direct contact with respiratory epithelium.^[60] Prolonged exposure often leads to epithelial hyperplasia, where secondary lamellae thicken due to cell proliferation, potentially increasing diffusion distance but serving as a barrier against irritants; for instance, in rainbow trout exposed to copper, hyperplasia can elevate lamellar thickness by 50%.^[61] In extreme pressure environments, such as deep-sea habitats, gill morphology adapts by reducing lamellar density to minimize structural stress and flow resistance. Deep-water species like the coelacanth (Latimeria chalumnae) exhibit sparsely distributed lamellae with a reduced overall surface area, featuring short filaments and a thick blood-water barrier (approximately 6 μm), which suits low-oxygen, high-pressure conditions at 100-400 m depths. This configuration supports lethargic lifestyles with minimal ventilation rates (3-4 movements per minute), prioritizing energy conservation over high-capacity gas exchange.^[62]

Evolutionary Development

The evolutionary origins of fish gills trace back to the pharyngeal slits of early chordates, which emerged approximately 500 million years ago during the Cambrian period.^[63] These structures, observed in fossils like the Middle Cambrian Pikaia gracilens, initially functioned primarily in filter-feeding, allowing water to pass through slits for capturing food particles while secondarily facilitating some gas exchange.^[63] Similarly, Cambrian vetulicolians exhibit evidence of pharyngeal slits and a pharynx, supporting the hypothesis that these openings evolved as a shared deuterostome innovation for both feeding and respiration in ancestral chordates.^[64] In early agnathans, such as ostracoderms from the Ordovician to Devonian periods, pharyngeal slits underwent a significant transition from dual-purpose organs to specialized respiratory structures. This shift occurred as feeding mechanisms evolved, with the development of rasping structures like dental plates in some ostracoderms, freeing the gills from primary filter-feeding duties and allowing them to optimize oxygen extraction from water.^[65] Common ancestors of lampreys and gnathostomes possessed fully developed gills capable of efficient respiration, indicating that this specialization predated the divergence of jawless and jawed vertebrates around 450 million years ago.^[66] A pivotal innovation in gnathostome evolution was the development of secondary lamellae on gill filaments, which dramatically increased surface area for gas exchange and enhanced respiratory efficiency in oxygenated aquatic environments. This adaptation likely arose in early jawed fishes during the Silurian-Devonian transition, enabling gnathostomes to outcompete agnathans by supporting higher metabolic demands associated with active predation. Fossil evidence from Devonian sarcopterygians, such as Eusthenopteron foordi, preserves distinct impressions of gill filaments and supporting rays, illustrating the structured complexity of these lamellated gills approximately 385 million years ago.^[67]

Pathologies and Interactions

Parasitic Infections

Fish gills are particularly vulnerable to parasitic infections due to their highly vascularized and delicate structure, which provides an ideal attachment site for ectoparasites.^[68] Common parasites targeting gills include monogeneans, copepods, and amoebae, which can impair respiratory function, cause tissue damage, and lead to secondary infections.^[69] Monogeneans, such as gill flukes of the genus Dactylogyrus, are flatworm ectoparasites that primarily infest the gills of freshwater fish like cyprinids.^[69] These parasites attach to gill filaments using specialized posterior attachment organs called opisthaptors, equipped with hooks that anchor into the epithelial tissue.^[70] This attachment causes mechanical damage, hyperplasia of the gill epithelium, excessive mucus production, and inflammation, which collectively reduce the gill's surface area for gas exchange.^[69] Heavy infestations often result in secondary bacterial or fungal infections on the damaged tissue, exacerbating mortality in affected fish.^[71] For instance, Dactylogyrus species can lead to significant respiratory distress and osmoregulatory dysfunction in infected hosts.^[72] Copepods, exemplified by species like Lernaea cyprinacea (anchor worm), are crustacean parasites that can attach to gill filaments in addition to skin and fins.^[73] After molting into their parasitic copepodid stage, these organisms embed into the host tissue using a holdfast structure, feeding on mucus, skin, and blood.^[74] Attachment on gills clogs the filaments and causes localized tissue necrosis, thereby reducing oxygen uptake efficiency and impairing overall respiration.^[75] In severe cases, the physical obstruction and blood loss from multiple attachments contribute to anemia and increased susceptibility to opportunistic pathogens.^[76] Amoebic gill disease (AGD) primarily affects salmonids in marine aquaculture and is caused by the protozoan ectoparasite Neoparamoeba perurans.^[77] The amoeba colonizes gill lamellae, inducing hyperplastic lesions, interlamellar fusion, and excessive mucus secretion, which severely compromise gill function and lead to respiratory failure if untreated.^[78] This pathology is characterized by raised white patches on the gills and can recur in infected populations, particularly under high stocking densities.^[79] Parasitic gill infections are highly prevalent in aquaculture settings, where intensive farming practices facilitate rapid parasite transmission and amplification.^[80] Economically, these infections cause substantial losses, with parasites estimated to account for 1-10% of global finfish production value, equating to $945 million to $9.45 billion annually, through direct mortality, reduced growth rates, and treatment costs.^[81] In regions like the UK, parasitic diseases contribute to 5.8-16.5% of annual aquaculture production losses across species.^[82] Control methods focus on breaking parasite life cycles and include freshwater baths, which effectively dislodge gill-attached parasites like monogeneans and N. perurans by osmotic shock, typically administered for 2-3 hours.^[83] Other approaches involve chemotherapeutants such as formalin or hydrogen peroxide baths, alongside biosecurity measures to prevent introduction from wild fish or equipment.^[80]

Gill Damage and Diseases

Bacterial infections represent a significant non-parasitic threat to fish gill health, with Flavobacterium columnare being a primary causative agent of columnaris disease. This Gram-negative bacterium adheres to gill tissues, leading to rapid proliferation and subsequent necrosis of the gill epithelium, which impairs respiratory and osmoregulatory functions. In infected fish, such as channel catfish and koi carp, the infection manifests as yellowish lesions and bacterial mats on the gills, often resulting in high mortality rates under stressful aquaculture conditions.^[84]^[85] Viral hemorrhagic septicemia (VHS), caused by the viral hemorrhagic septicemia virus (VHSV), particularly affects the gill epithelium in freshwater species like rainbow trout. The virus targets epithelial cells, inducing hemorrhagic lesions and epithelial sloughing that compromise gas exchange and ion regulation. Freshwater isolates of VHSV demonstrate higher virulence in infecting gill cells compared to marine strains, correlating with increased mortality in susceptible populations.^[86]^[87] Physical damage to fish gills arises from mechanical abrasion, such as contact with fishing nets or suspended sediments, and environmental stressors like low dissolved oxygen levels. Abrasion from nets or abrasive particles causes direct injury to delicate gill filaments, triggering inflammatory responses including hyperplasia, where excessive cell proliferation thickens the lamellae and reduces oxygen diffusion efficiency. Similarly, hypoxia from low dissolved oxygen prompts adaptive hyperplasia as a compensatory mechanism, but severe cases lead to lamellar fusion and impaired ventilation, exacerbating mortality during fish kills.^[88]^[89]^[90] Toxic exposures to heavy metals, notably copper, induce gill damage by disrupting ionoregulatory processes in chloride cells. Copper ions accumulate primarily in the gills, inhibiting key enzymes such as Na⁺/K⁺-ATPase, which is essential for active ion transport and osmotic balance. This enzymatic inhibition leads to ionoregulatory failure, cellular necrosis, and hyperplasia in chloride cells, particularly in freshwater fish exposed to elevated copper concentrations from industrial effluents or aquaculture treatments.^[91]^[92]

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[12]
Fick's Diffusion Laws and Scaling of the Gill Surface Area ... - MDPI
The oxygen consumption of adult fish (Q) is proportional to their body weight (W) raised to a power, estimated as the slope (dQ) of a linear regression of ...
[13]
Fine structure of the respiratory lamellae of teleostean gills
This layer of cytoplasm is formed by flange-like extensions of the pillar cells. The resulting location of the pillar cell perikarya between the surfaces of the ...
[14]
Contractile filamentous material in the pillar cells of fish gills
Sep 1, 1973 · The secondary lamellae of the gill filaments are the sites of gas exchange in fish gills. They are mainly composed of 2 epithelial sheets ...
[15]
WFS 550 Fish Physiology- Respiratory Pump
In most fish, the system is unidirectional; water always moves through the mouth and one way across the gills, then out through the operculum.Missing: mechanism | Show results with:mechanism
[16]
https://link.springer.com/article/10.1007/BF02044316
[17]
Oxygen utilization and the branchial pressure gradient during ram ...
Jan 1, 2012 · Lamnids and tunas are also obligate ram ventilators, meaning they depend on continuous swimming to force water through the gills.
[18]
The Multifunctional Fish Gill: Dominant Site of Gas Exchange, Osmoregulation, Acid-Base Regulation, and Excretion of Nitrogenous Waste | Physiological Reviews | American Physiological Society
Below is a merged summary of the passive diffusion of O2 and CO2 in fish gills, consolidating all information from the provided segments into a comprehensive response. To maximize detail and clarity, I’ve organized key information into a table in CSV format, followed by a narrative summary that integrates additional details not suited for the table. The response retains all mentioned data, including citations, URLs, and specific references, while avoiding redundancy where possible.
[19]
Henry's Law - StatPearls - NCBI Bookshelf - NIH
This law defines the relationship between the partial pressure of gases overlying a solution and the gases' ability to dissolve in that solution.
[20]
Ion regulation in fish gills: recent progress in the cellular and ...
Gills are one of the major organs conducting the internal ionic and acid-base regulation, with specialized ionocytes as the major cells carrying out active ...
[21]
Fish gills: mechanisms of salt transfer in fresh water and sea water
Functional studies show that the gill is responsible for the net absorption of Na+ and Cl- occurring in fresh water and extrusion of these ions in sea water.
[22]
The Dimensions of Fish Gills in Relation to Their Function
It is concluded that more active fish not only have larger gill areas but that the conditions for gaseous exchange are better than for more sluggish forms and ...
[23]
Structural and functional studies of the king salmon, Oncorhynchus ...
Cathodal hemoglobin components exhibited a higher oxygen affinity (P50 = 10.2 mm at 13 degrees C, pH 7.3) than the anodal hemoglobin components (P50 = 21.8 mmHg ...Missing: values | Show results with:values
[24]
Modulation of Allosteric Control and Evolution of Hemoglobin - PMC
Mar 22, 2023 · The peculiar O2 binding properties of fish hemoglobins has solicited over the years the curiosity of many blood physiologists and biochemists.<|control11|><|separator|>
[25]
A broad diversity in oxygen affinity to haemoglobin - PMC - NIH
Oct 9, 2020 · Standard p50 in humans is 26.9 mmHg at pH 7.4 and 37°C. Increased values of temperature, hydrogen ions (acidosis), pCO2, and 2,3-BPG all lower ...
[26]
Red blood cell potentiation of hemoglobin-oxygen unloading in fish
The Bohr effect was discovered over a century ago (4) and describes how metabolically produced carbon dioxide (CO2) acidifies the blood and reduces the affinity ...
[27]
Bohr Effect - an overview | ScienceDirect Topics
The Bohr effect refers to the physiological phenomenon where an increase in carbon dioxide concentration or a decrease in pH reduces hemoglobin's affinity ...
[28]
Root Effect Haemoglobins in Fish May Greatly Enhance General ...
Oct 5, 2015 · Root effect Hbs can enhance O 2 release to the tissues by 73.5% in trout; whereas, the Bohr effect alone is responsible for enhancing O 2 release by only 1.3% ...
[29]
Gill Blood Flow Control - ScienceDirect.com
In teleost fish, blood from the ventral aorta enters the gills via four pairs of afferent ... The sphincter of the efferent filament artery in teleost gills.Missing: scholarly | Show results with:scholarly
[30]
https://www.necropsymanual.net/en/teleosts-anatomy/respiratory-system/
[31]
https://www.sciencedirect.com/topics/veterinary-science-and-veterinary-medicine/buccal-pumping
[32]
Respiratory system - Fish Necropsy Manual
Teleosts have four holobranches and one hemibranch on each side. Each holobranch is supported by a gill arch and is made of two divergent hemibranches that ...Missing: osteichthyes | Show results with:osteichthyes
[33]
Buccal Pumping - an overview | ScienceDirect Topics
The vast majority of fish species use buccal pumping for gill ventilation, whereby a current of water is drawn into the mouth and over the gills via actively ...
[34]
Mechanics of Respiratory Pumps - ResearchGate
Aug 7, 2025 · The gills are protected by a bony operculum that also acts (in conjunction with a backflow valve in the mouth) as a pump to move water ...
[35]
Cardiorespiratory modifications, and limitations, in post-smolt growth ...
Apr 1, 2006 · ... salmon are compensating for increased metabolic demands using a `whole system' approach. Interestingly, however, our results do not support ...
[36]
Fish gill surface area can keep pace with metabolic oxygen ...
A similar lamellar density across acclimation temperature in the present study may therefore aid in explaining why MMR was unable to increase further at 19 • C, ...
[37]
The chloride cell: structure and function in the gills of freshwater fishes
Chloride cells are the main site for Ca2+ and Cl- influx, and perform an integral role in acid-base regulation, with surface area changes during alkalosis and ...Missing: bony | Show results with:bony
[38]
Blue in the Face: Breathing Underwater (U.S. National Park Service)
Jan 24, 2020 · In contrast, bony fish have a special bone called the operculum that lets them use their gills without ram ventilation. The operculum rests ...Missing: teleosts | Show results with:teleosts
[39]
The Origin of Vertebrate Gills - PMC - NIH
Feb 9, 2017 · It is thought that gills evolved independently in cyclostomes (jawless vertebrates—lampreys and hagfish) and gnathostomes (jawed vertebrates— ...
[40]
The Lamprey: A jawless vertebrate model system for examining ...
Their pharynx is perforated by seven round gill slits, which open into muscular pharyngeal pouches. They lack paired fins, but have dorsal, caudal, and anal ...
[41]
Feeding and Breathing in Lampreys - Karger Publishers
Jan 8, 2008 · Ammocoete larvae use a velar pump for unidirectional ventilation and suspension feeding. In both lampreys and ammocoetes the branchial basket ...
[42]
It's all in the gills: evaluation of O2 uptake in Pacific hagfish refutes a ...
Hagfish primarily relied on the gills for both O2 uptake (81.0%) and ammonia excretion (70.7%). Following exercise, both O2 uptake and ammonia excretion ...Missing: efficiency | Show results with:efficiency
[43]
Mitochondria from the systemic heart of Pacific hagfish (Eptatretus ...
The respiration rate of mitochondria in permeabilized hagfish heart tissue is comparatively low compared to that of other ectothermic species.
[44]
The hagfish genome and the evolution of vertebrates - Nature
Jan 23, 2024 · As the only surviving lineages of jawless fishes, hagfishes and lampreys provide a crucial window into early vertebrate evolution.
[45]
Distinct freshwater and seawater isoforms of Na+/K+-ATPase in gill ...
Abstract. Gill Na(+)/K(+)-ATPase (NKA) in teleost fishes is involved in ion regulation in both freshwater and seawater.
[46]
Differential regulation of select osmoregulatory genes and Na+/K+ ...
Integral to the osmoregulation process in fish is Na+/K+-ATPase (NKA), an ion pump that powers ion transport across the membrane and is abundant in the gill.
[47]
Ammonia Production, Excretion, Toxicity, and Defense in Fish
Therefore, a significant portion of ammonia can be excreted through NH 4 + diffusion through the paracellular route in seawater fishes (Goldstein et al., 1982).
[48]
Mechanisms of Ammonia Excretion Across Fish Gills - ScienceDirect
In fresh water, ammonia excretion likely takes place via passive NH 3 diffusion down favourable blood-gill water P NH3 gradients.
[49]
Ammonia distribution and excretion in fish - PubMed
The mechanisms of branchial ammonia excretion vary between different species of fish and different environments, and primarily involves NH3 passive diffusion ...
[50]
Gill Morphology and Acid-Base Regulation in Freshwater Fishes
This review examines the recent advances in our understanding of the mechanisms of ion transport and acid-base regulation in the freshwater fish gill.
[51]
New insights into the many functions of carbonic anhydrase in fish gills
Acid–base regulation in fish relies primarily on modulation of Cl−/HCO3− and Na+/H+ exchanges at the gill so as to adjust plasma HCO3− concentration (Claiborne ...
[52]
Rapid blood acid–base regulation by European sea bass ... - NIH
Fish have a great capacity to restore blood pH to compensate for CO2-induced respiratory acidosis, which is largely achieved by excreting excess H+ and ...
[53]
Developing zebrafish utilize taste-signaling pathways for oxygen ...
Sep 23, 2024 · We show that Merkel-like cells (MLCs), which are part of the taste-bud complex, function as O 2 chemoreceptor cells in larval zebrafish.
[54]
Expression of taste receptors in Solitary Chemosensory Cells of ...
In fish, the SCCs appear to share some amino acid receptors with taste buds [29]. In the last decade, morphologically similar cells were described in the nasal ...
[55]
Nervous control of the gills - ScienceDirect.com
From here, blood enters the gill arches via the afferent branchial artery, and the gill filaments via the afferent filament arteries. Blood flow continues ...Missing: scholarly | Show results with:scholarly<|separator|>
[56]
Chronic hypoxia and Cu2+ exposure induce gill remodeling of ...
Fishes are capable of remodeling gills to adapt to hypoxia by reducing the height of interlamellar cell mass (ILCM) and expose the gill lamellar under hypoxia ...
[57]
Gill remodeling in fish – a new fashion or an ancient secret?
Jul 15, 2007 · Gill remodeling is when fish reversibly change gill morphology, like in crucian carp, by removing cell mass to expose more respiratory surface ...
[58]
Measuring oxygen uptake in fishes with bimodal respiration - Lefevre
Sep 11, 2015 · Air breathing is believed to have evolved in response to aquatic hypoxia and, probably, associated hypercapnia. The review ends by considering ...
[59]
Do air-breathing fish suffer branchial oxygen loss in hypoxic water?
Sep 13, 2023 · Many teleost fish resort to air-breathing in hypoxic and warm tropical waters and this ability has evolved independently more than 80 times [1].
[60]
Physiological mechanisms used by fish to cope with salinity stress
Jun 1, 2015 · In addition, euryhaline fish achieve a switch from plasma hyper- to hypo-osmoregulation by increasing cell proliferation and turnover and via ...
[61]
Changes in osmotic water permeability of the eel gills during ...
The gradual increase in the gill water permeability during seawater transfer is correlated with an increase in the number of chloride cells. In scanning ...Missing: proliferation | Show results with:proliferation
[62]
Fish Gill Structural Changes Induced by Toxicants and Other Irritants
Here I quantitatively review the literature on how fish gill morphology is affected by chemical and physical irritants in the surrounding water.
[63]
Fish Gill Structural Changes Induced by Toxicants and Other Irritants
Aug 6, 2025 · Acute heavy metal toxicity to fish has been attributed to the coagulation or precipitation of mucus on the gills and/or to cytological damage ...
[64]
[PDF] The long-time adaptation of coelacanths to moderate deep water
May 14, 2019 · In adult specimens of L. chalumnae gills are poorly developed, like in deep-water fishes, with a reduced surface area and sparse lamellae that ...
[65]
The Middle Cambrian fossil Pikaia and the evolution of chordate ...
Jun 13, 2012 · Pharyngeal pores or slits, where they occur, are assumed to play an ancestral role in deposit- or filter-feeding as a means for disposing of ...
[66]
Evidence for gill slits and a pharynx in Cambrian vetulicolians
Oct 2, 2012 · ... most seminal was the evolution of openings that define the pharyngeal gill slits of hemichordates (and some extinct echinoderms) and chordates.
[67]
The Origin Of Vertebrates And The Rise Of Fishes
It also allowed gills to specialize in gas exchange (rather than serve the dual function of gas exchange and filter feeding). 2.Missing: transition | Show results with:transition
[68]
Early vertebrate evolution: pharyngeal structure and the origin of ...
Aug 7, 2025 · It is concluded that the common ancestors of gnathostomes and lampreys were not simple animals, but fish with fully developed gills that could ...
[69]
[PDF] SPECIALIZATIONS IN EARLY VERTEBRATES
distinct impressions of the gills and their supporting gill-rays are preserved in one specimen of the Devonian osteolepiform fish Eusthenopteron (fig. 5 D) ...
[70]
FA28/FA033: Monogenean Parasites of Fish - University of Florida
Secondary infections with bacteria and water molds are common on tissue that has been damaged by monogeneans. In marine fishes, the capsalid monogeneans may ...Missing: Dactylogyrus | Show results with:Dactylogyrus
[71]
Dactylogyridae) on Gills of a Model Fish, Gobiocypris rarus ...
Jan 28, 2023 · Dactylogyrids can infect the gills of cypriniform fishes [18], causing serious hyperplasia of the gill filament epithelium, copious mucus, and ...
[72]
12. monogenea.
Monogeneans are flatworms (Platyhelminthes), ectoparasitic and attached by special posteriorly positioned attachment organs to their host's skin or gills.
[73]
[PDF] Introduction to Freshwater Fish Parasites 1 | RWFM Extension
Secondary infection by bacteria and fungus is common on tissue with monogenean damage. Gyrodactylus and Dactylogyrus are the two most common genera of ...
[74]
Parasite Diversity in a Freshwater Ecosystem - PMC - NIH
Jul 29, 2023 · Monogeneans cause an increase in fish mortality due to various infections such as respiratory problems, anemia, and osmoregulatory dysfunction, ...
[75]
Lernaea (Anchorworm) Infestations in Fish - University of Florida
At that point they molt into the first copepodid stage, become parasitic, and attach to a host, often on the gills. Over the next 7 days, the parasite ...
[76]
Copepods - Fish Pathogens
Parasitic copepods feed on skin, mucus and blood from their hosts. Usually there are few parasites present on fish and they go unnoticed, however occasionally ...
[77]
Natural infestation of an anchor worm, Lernaea sp. in cage culture of ...
Lernaea sp. infestation can occur in skin, fins, gills, and oral cavity but in the present outbreak, it was observed only on the skin as the fish juvenile were ...Missing: uptake | Show results with:uptake
[78]
[PDF] Diseases of Wild and Cultured Fishes in Alaska
Epidermal and gill tissues may become necrotic with secondary infections by fungi and bacteria. These parasites are found worldwide. II. Host Species. A ...
[79]
Host-Parasite Interaction: Salmon & Neoparamoeba perurans
Marine farmed Atlantic salmon (Salmo salar) are susceptible to recurrent amoebic gill disease (AGD) caused by the ectoparasite Neoparamoeba perurans over ...
[80]
Mucosal responses in the skin and gill filaments of Atlantic salmon ...
Oct 31, 2024 · Amoebic gill disease (AGD), primarily affecting salmonids, is caused by Neoparamoeba perurans, which colonizes the fish gills and is a ...
[81]
In vitro gill cell monolayer successfully reproduces in vivo Atlantic ...
An in vitro model to study the host response to Neoparamoeba perurans, the causative agent of amoebic gill disease (AGD), was evaluated.
[82]
Control of parasitic diseases in aquaculture - PMC - NIH
The most direct way to limit development of a parasite infection in a host population is to block the life cycle. For certain parasites with free-living ...
[83]
Economic impacts of aquatic parasites on global finfish production
Assuming parasites are responsible for losses of 1% to 10% of harvest-size fish, the value of these fish was estimated at $945.00 million to $9.45 billion.
[84]
An overall estimation of losses caused by diseases in the Brazilian ...
Jul 7, 2017 · Current studies suggest that parasites account for an annual loss of 5.8–16.5% of UK aquaculture production considered all the species in both ...
[85]
Evaluation of the Infectious Potential of Neoparamoeba perurans ...
Freshwater bathing for 2–3 h is the main treatment to control amoebic gill disease of marine-farmed Atlantic salmon. Recent in vitro studies have demonstrated ...
[86]
Columnaris disease in fish: a review with emphasis on bacterium ...
columnare infections may result in skin lesions, fin erosion and gill necrosis, with a high degree of mortality, leading to severe economic losses. Especially ...
[87]
Flavobacterium columnare (Flexibacter columnaris) associated with ...
Flavobacterium columnare (Flexibacter columnaris) associated with severe gill necrosis in koi carp (Cyprinus carpio L)
[88]
Differences in virulence of marine and freshwater isolates of viral ...
Differences in virulence of marine and freshwater isolates of viral hemorrhagic septicemia virus in vivo correlate with in vitro ability to infect gill ...
[89]
The Viral Hemorrhagic Septicemia Virus (VHSV) Markers of ... - NIH
Oct 20, 2020 · Viral hemorrhagic septicemia virus (VHSV) is a highly contagious virus leading to high mortality in a large panel of freshwater and marine fish ...
[90]
[PDF] Essential Fish Habitat (EFH) Assessment New Bedford/Fairhaven ...
Gill nets can damage fish either via compressing their gills leading to suffocation or via gill injury while struggling in the net (WADFW, 1997). For ...
[91]
[PDF] Effects of Turbidity and Suspended Solids on Salmonids - WSdot.com
Fish gills are delicate and easily damaged by abrasive silt particles. As sediment begins to accumulate in the filaments, fish excessively open and close their ...
[92]
[PDF] Causal Analysis of Fish Kills in the Shenandoah and Potomac Rivers
May 2, 2007 · Upon examination, affected fish had gill hyperplasia (thickening of the gill filaments) that diminished the surface area for oxygen exchange.
[93]
Review of Copper and Copper Nanoparticle Toxicity in Fish - PMC
Based on the toxicity experiments conducted with diverse fish species, waterborne copper exposure can induce various kinds of organ damage in the gills, liver, ...
[94]
[PDF] Aquatic Life Ambient Freshwater Quality Criteria for Copper
Feb 14, 2007 · ... copper toxicity therefore could be due to effects on copper bioavailability. ... fish gill). For certain ligands, some studies have.