Alsafy et al.

BMC Zoology (2025) 10:3 BMC Zoology

https://doi.org/10.1186/s40850-025-00223-5

RESEARCH Open Access

Anatomy, histology, and morphology

of fish gills in relation to feeding
habits: a comparative review of marine
and freshwater species
Mohamed A. M. Alsafy1* , Hanan H. Abd-Elhafeez2, Ahmed M. Rashwan3,4, Atef Erasha5, Safwat Ali6 and Samir A.
A. El-Gendy1

Abstract
This systematic review highlights the similarities and variations in gill morphology, histology, and anatomical
structure between differing fish species. The gill system consists of mainly four pairs of gill arches in most teleost
fishes, such as sea bass, sea bream, grouper, and red porgy, etc., while it consists of three pairs of gill arches in
pufferfish and striped-red mullet fish. However, Clarias gariepinus had five pairs, including an additional rudimentary
fifth-gill arch. The gill structure consisted of gill arches, gill rakers, gill filaments, and secondary lamellae with varied
shapes of gill arches such as hook, semilunar, L-shapes, and crescentic shapes. Each gill arch carried mainly two
rows of gill rakers, lateral and medial, present in most teleost fishes (Mugil cephalus, Boops boops, Pagrus pagrus,
Sparus aurata, European hake, Puffer fish, grey gurnard, sea bass, and sea bream). An additional row appears in
Clarias gariepinus or two rows (accessory) in dusky grouper fish. The length and shape of gill rakers are mainly
related to feeding habits. The gill rakers in lateral rows are longer, equal, or more in number and more developed
than those of the medial rows, except at three gill arches in striped-red mullet fish, the second and third gill
arches in pufferfish, and the fourth arch in Pagrus pagrus. gill rakers are absent at the first and second gill arches
in Bagrus bayad. The gill arch carries additional structures, such as the air-breathing dendritic organ of the catfish,
located in the suprabranchial chamber caudodorsal to the gills and composed of two main parts: small and large
ones originated by main stems from the second and fourth-gill arches, respectively. The interbranchial septum
can be smooth, form a median crest (seabream), or carry teeth or spines (seabass, pufferfish). Four transversely
raised areas on each side are connected by transverse lines caudal to the base of the tongue (Bagrus bayad) and
an elevated part at the level of the third-gill arch (Tilapia zilli). Scanning electron microscopy explained the micro-
anatomical structures as varied shapes of pavement cells, mucus cell openings, taste buds on the gill arch, varied
shapes of grooves or structures and spines near the gill filament side, varied shapes of gill rakers and their spines,
and heights in varied feeding types of fish. Histological findings revealed various types of cells, such as superficial
pavement cells, large chloride cells, mucous goblet cells, and basal epithelial cells. The lymph space is situated

*Correspondence:
Mohamed A. M. Alsafy
[email protected]; [email protected]
Full list of author information is available at the end of the article

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Alsafy et al. BMC Zoology (2025) 10:3 Page 2 of 21

within the gill arch epithelia and is encompassed by cells that resemble tenocytes. The lymph space contains
many types of immunological cells, including lymphocytes, granular leukocytes, and rodlet cells. The gill arch
comprises sensory structures known as neuromasts and hyaline cartilaginous support. This review underscores the
intricate relationship between gill structure and feeding habits across marine and freshwater fishes, highlighting
the importance of understanding these variations for ecological, evolutionary, and aquacultural applications and
feeding habits.
Keywords Fish, Gill, Gill arch, Gill raker, Gill filaments

Background scanning electron microscopy [33–36] and light micros-

The gills form a highly characteristic feature of fishes, and copy slides with PAS stain, and semithin sections with
their presence has a marked effect on the anatomy and toluidine blue stain, techniques according to [20, 37–41].
functioning of the rest of the animal [1]. The gills play a
significant role in adapting the fish to their environment. Anatomy of gills
The primary functions of gills are gas exchange and waste The gill system was confined within two interconnected
elimination, and the gill epithelium also plays an osmo- gill chambers. The gill chambers were bounded ventrally
regulatory role [2–7]. Typically, gills are the gas exchange by the mandible, dorsally by the roof of the oral and pha-
organs of water-breathing fishes, but in some species, ryngeal cavities, and laterally by the operculum. While
they are also involved in gas exchange with the air [1, 8, medially, they were continuous with each other. Most
9]. gills of the teleost fishes’ were arranged on each side from
The gills, like other systems in the fish body, undergo lateral to medial as the first, second, third, and fourth
numerous modifications to adapt to the specific physico- gills [3, 4, 17]. In the dusky grouper [19], each gill was
chemical conditions under which they must function [1]. semilunar in shape, consisting of a gill arch that carried
There is a notable adaptation of fish to their environment gill rakers on its concave border and gill filaments on its
and feeding habits, which is reflected in the morpho- convex border (Fig. 1A and B).
logical characteristics of their gills [4, 10–13]. There are Sea bream and sea bass are carnivorous fish that pri-
many differences in the structural components of the gill marily feed on large-sized food items such as mollusks,
arches, gill filaments, and gill rakers [3, 4, 12–14]. small fish, crustaceans, insects, and decaying organic
The intricacies of gill morphology have captivated matter. Consequently, their gills are characterized by a
researchers, prompting numerous studies that delve into prominent angle of curvature at the union of short epi-
the gill structures of various fish species [2–4, 9, 15–23]. branchials and long ceratobranchials from the first to
The organization of gill filaments and gill rakers is closely fourth pairs of gill arches toward the dorsal side, simi-
related to the feeding habits of the fish [24–26]. The lar to the carnivorous feeder catfish Rita rita [42] and
size and number of gill rakers control the size of food Eugerres brasilianus, as well as the omnivorous feeder
ingested; fish species with many and long gill rakers are [31]. In contrast, gill arches in filter-feeding mullets, such
typically filter feeders, whereas those with few and short as Mugil cephalus, show a lack of curvature or display an
gill rakers tend to be omnivores or carnivores [20, 27, 28]. acute angle of curvature in the middle of the gill arches,
The epithelial cells covering the gill arches, gill rak- indicating the degree of pharynx expansion [43].
ers, and gill filaments exhibit variable surface features in
previously studied fish species, including the presence of Gill arch
micro ridges forming specific patterns, a mosaic of pave- Each gill arch had two extremities: rostral and caudal.
ment cells, varied sizes of mucous and chloride cell pores, The rostral extremities of the gill arches were united,
numerous pointed spines, and various types of taste buds forming an interbranchial septum between the contralat-
[3, 4, 17, 29–31] eral gills. The septum exhibited a median elevation like
a crest in sea bream (Fig. 2A, B, C). At the same time, it
Material and methods appeared flattened dorsoventrally. It carried two rough,
A systematic review of the literature was conducted spiny small gill rakers at the level of the third gill in sea
using three databases (Nusearch, CAB Abstracts, and bass (Fig. 3A, B). The gills on both sides diverged caudol-
PubMed), as well as ResearchGate, Google Scholar, and aterally, leaving a triangular area bounded rostrolaterally
general internet searches. A lot of relevant papers were by the fourth pair of gills and occupied by the floor of the
collected and evaluated for this review [32]. pharynx (Figs. 2B, C, 3A, B)—the caudal extremities of
We selected figures from our unpublished data. We the four-gill arches curved dorsally, rostrally, and slightly
collected fish from the Mediterranean Sea, the Red Sea, ventrally. The gill arches were connected and attached
and the Nile River. We prepared gross anatomy figures, to the operculum’s medial surface and the pharynx’s
Alsafy et al. BMC Zoology (2025) 10:3 Page 3 of 21

Fig. 1 Gross images of the grouper fish: A rostral view of the opened mouth and B lateral view of the gills with reflected operculum that explained the
structures and position of the gills. (A1, A2, A3, and A4) 1st–4th gill arches. rackers (R), a lateral row of gill rakers (LR), esophagus (O), interbranchial septum
(IS), the mandible (M), the roof of the pharynx (RP), the roof of the oral cavity (RO), pharyngeal teeth (PT), chewing pads (Cp), operculum (OP), rostral
extremities (RE), caudal extremities (CE), short epibranchial (eb), long ceratobranchial (cb), angle of epi-ceratobranchial union (ag), gill raker (R), gill arch
(A), gill filaments (F), and tongue (T)

Fig. 2 Gross images of sea bream fish: A lateral view of the fish and B and C the oropharyngeal cavity explained the following structures. (A1, A2, A3,
and A4) 1st–4th gill arches, lateral row of gill rakers (LR), interbranchial septum (IS), roof of pharynx (RP), roof of the oral cavity (RO), pharyngeal teeth (PT),
median crest (MC), chewing pads (Cp), tongue (T), and apical pouch (AP)
Alsafy et al. BMC Zoology (2025) 10:3 Page 4 of 21

Fig. 3 A, A*, B and B* Gross images and C and D scanning electron micrographs of the gills of the Sea bass fish. A Lateral view of the gills after removing
the operculum. B Rostral view to the gills chamber. C, and D SEM images of the 1st gill arch. (A1, A2, A3, and A4) 1st–4th gill arches, long gill rakers (R1),
short gill rakers (R2), angle of epi-ceratobranchial union (ag), interbranchial septum (IS), median crest (MC), pharyngeal teeth (PT), pavement cells (PC),
pores of mucous cell opening (P), tongue (T), spines (S), microplicae (MP), and taste buds (TB)

dorsolateral wall. The length and gaps between the four arches appeared curved dorsally. They were connected
gill arches decreased medially, while the width and thick- and attached to the dorsolateral wall of the pharynx. The
ness of the four gill arches were similar. The gill arch was lengths of the three gill arches were measured at 4.7 cm,
divided into a long ceratobranchial and a short epibran- 4.5 cm, and 4.1 cm, respectively, from lateral to medial.
chial. The length ratio of the two parts varied from one Each gill arch was divided into a long ceratobranchial
gill arch to another. The union of the two parts of the gill and a short epibranchial part, demarcated by a slightly
arches toward the dorsal side showed a more prominent prominent angle. The length ratio of the two parts varied
angle of curvature in sea bass than in sea bream [4]. from one gill arch to another, decreasing from lateral to
The gill arch plays a crucial role in immunity [44]. The medial.
gill arches can effectively buffer the water flow pressure. In grey gurnard fish [15], the gill arch had a crescen-
The circular microridges can fix the mucus on the sur- tic shape. The rostral extremities of the gill arches united
face, and the strength of cells can be enhanced by the to form a wide interbranchial septum that appeared as a
structure of circular microridges that can alleviate the quadrilateral narrow structure carrying four transverse
mechanical damage of water flow. The gill arch taste buds elevated crests.
have a role in food selectivity [19]. In the striped-red mullet [15], the interbranchial sep-
In the dusky grouper [19], the gill chamber had four tum had a median longitudinal elevated crest. The gill
gills on each side. They were semilunar in shape. The arches were connected and attached to the dorsolateral
length of the gill arches from the 1st to the 4th gill arch wall of the pharynx. Each gill arch lacked a distinct angle
were 5.27 cm, 4.2 cm, 3.2 cm, and 2.8 cm, respectively. between the ceratobranchial and epibranchial parts,
Each gill arch had a long horizontal part. The ceratobran- making it difficult to differentiate between them. The gill
chial parts were four times the length of the short epi- arches were connected rostrally and caudally.
branchials, and the angle between the two parts of the gill In European hake [17], the thickness and width of the
was around 78° (Fig. 4A, B, C). four-gill arches were similar, while the length and gaps
In Puffer fish [45], the gill arches had crescentic shapes between the gill arches decreased medially. The lengths
and were directed rostroventrally. The rostral border of of the four gill arches were measured at 5.3 cm, 4.7 cm,
the gill arches formed a wide interbranchial septum that 4.5 cm, and 4 cm, respectively. Each gill arch had a long
appeared quadrilateral and flattened dorsoventrally, car- ceratobranchial part and a short epibranchial part. The
rying a line of small spiny gill rakers ventromedially to length ratio of the two parts varied between the gill
the third-gill arch. The caudal extremities of the three gill
Alsafy et al. BMC Zoology (2025) 10:3 Page 5 of 21

Fig. 4 A, B, and C Gross images and D and E scanning electron micrographs of the gills of the dusky grouper fish. A Lateral view image of the dusky
grouper fish. B Rostral view to the gills chamber. C Lateral view of the 1st and 2nd gill arch. D and E SEM images of the medial and dorsal view of the 1st
gill arch. (A1, A2, A3, and A4) 1st–4th gill arches, long gill rakers (R1), short gill rakers (R2), accessory gill rakers (Ra), angle of epi-ceratobranchial union (ag),
spines (S), solitary spines (So), pavement cells (PC), esophagus (E), microplicae (M), and gill filaments (F)

arches, with both parts demarcated by an angle, which consisting of four transverse raised areas on each side,
was very narrow at the level of the second-gill arch. connected by transverse lines caudal to the base of the
In Bagrus bayad [3], the gill arch has two parts: a long tongue. The lengths of the gill arches were 7 cm, 5.5 cm,
ceratobranchial and a short epibranchial, clearly forming 5.5 cm, and 5 cm for the 1st, 2nd, 3rd, and 4th gill arches,
an angle between them in the first three gill arches, but respectively.
not obvious in the fourth-gill arch. The gill arch was cres- In the Nile River: Oreochromis niloticus, Chrysichthys
centic in shape and carried the gill rakers on the rostral auratus, and Clarias gariepinus [29], the gill system con-
concave border. The gill arches were attached rostrally sisted of four pairs of gills. Clarias gariepinus also had a
to the mandible, forming a wide interbranchial septum rudimentary fifth gill. Each gill was semilunar in shape,
Alsafy et al. BMC Zoology (2025) 10:3 Page 6 of 21

and the interbranchial septum was flattened dorsoven- water flow initially and then acted as a sieve, filtering
trally. The gills on both sides diverged caudolaterally, food particles by trapping them with their mucous cover
leaving a triangular-shaped area bounded rostrolaterally before ingestion [53].
by the fourth pair of gills and occupied by the floor of the Gill rakers are used to capture food and facilitate feed-
pharynx. This floor was modified into two distinct struc- ing behavior. Gill rakers varied in size and number among
tures: the hypopharyngeal bone carrying the pharyngeal fish that consumed large prey. Plankton feeders possessed
teeth and the lower pharyngeal jaw. The roof of the phar- elongated, numerous, and diverse lamellae or ornamen-
ynx, opposite the lower pharyngeal jaw, was modified tal gill rakers that functioned as effective sieves, allowing
into an oval-shaped structure. The gaps between the gill water to pass while trapping solid food particles [54, 55].
arches were wider in Clarias gariepinus than in the other The number, shape, and spacing of gill rakers reflected
two species and narrower in Oreochromis niloticus. the feeding behavior of fish species; fewer short gill rak-
In Clarias gariepinus [29, 46, 47], the breathing den- ers were found in carnivores and omnivores, whereas
dritic organ situated in the suprabranchial chamber cau- filter feeders exhibited numerous long gill rakers [4, 19,
dodorsal to the gills. This organ consisted of small and 52, 56–58]. Gill rakers are presented in small numbers in
large parts originating from the main stems of the sec- fish that consume large meals. Fish that only eat plankton
ond and fourth-gill arches. The size of the rostral small have long and varied lamellae, or ornate gill rakers, that
part was nearly half that of the large one, occupying the capture solid nutritional grains while allowing water to
majority of the rostral compartment of the suprabran- pass through during respiration [59]. Gill rakers are not
chial chamber. The large caudal part occupied most of suggested as a physical colander or strainer during filtra-
the middle caudal compartments of the suprabranchial tion. However, it should be noted that the flow dynam-
chamber. Both parts were connected to their corre- ics of liquids generate numerous vortices in the liquids
sponding gill arches by a cartilaginous joint. They origi- received into the mouth, resulting in cross-flow filtration.
nated from a small smooth surface main stem, divided The distance between the gill rakers is critical in remov-
into several secondary branches, ending in bulbous-like ing particles from the suspension [60]. Furthermore, gill
structures. rakers are the anatomical part investigated in the discov-
In Tilapia zilli [48], the gill system is located within ery of new of new species [61], and they serve as a model
two connected gill chambers. It was bounded by the for the industrial design of commercial filtration systems
mandible (ventrally), the operculum (laterally), the roof [60].
of the oral cavity (dorsally), and the base of the pectoral In carnivorous fish, the lateral gill rakers of the first-
fin (caudally). The gill arches were crescentic in shape, gill arch often bore small spines to prevent the escape of
carrying a row of gill filaments on their convex border slippery prey, whereas subsequent rows had shorter gill
and two rows of gill rakers on their concave border. The rakers [4, 53, 62]. Anterior gill rakers on the first gill arch
interbranchial septum was wide at the rostral part, quad- appeared as clusters of small tooth patches adapted for
rilateral in shape, with an elevated portion at the level of piscivorous feeding, while in herbivorous feeders, they
the third-gill arch. An angle in each gill arch divided it resembled needle-like spines with secondary projections
into a dorsal long ceratobranchial and a ventral short epi- (Fig. 3A, A⁎). Detritivores and planktivores typically pos-
branchial part. sessed the highest number and length of anterior gill rak-
In common carp (Cyprinus carpio), the interbranchial ers [63, 64].
septum connects the gills on both sides. The common European whitefish with sparse gill rakers tended to
carp’s gill has four gills on each side. It was arranged in consume mollusks, crustaceans, and insect larvae, while
a cranial-caudal direction. Each gill arch had two lat- those with denser gill rakers fed on zooplankton, chiron-
eral and medial rows of gill rakers; the length of the gill omid pupae, and surface insects [63].
arch decreased from the right first to the fourth gill arch In species like sea bass, two types of gill rakers were
(26.02 ± 2.04 mm, 24.69 ± 2.08 mm, 22.47 ± 1.19 mm, observed: well-developed gill rakers with minute spines
19.35 ± 11.11 mm) [49]. on the first gill arch and shorter gill rakers on subsequent
gill arches [53, 62]. These gill rakers varied in length and
Gill rakers structure across the gill arches, with sea bream display-
Gill rakers were cartilaginous or bony projections located ing nearly parallel arrangements and sea bass exhibiting
on the gill arches that aid food collection and feeding interdigitated patterns [4]. The gill rakers in sea bream
habits [20]. their structure and shapes varying accord- were short and wide-based with sharp ends, whereas in
ing to the feeding habits of the fish [50] (Fig. 4B, C). sea bass, they were cylindrical with tapering ends and
They were distributed unevenly across the branchial gill medial surfaces resembling saw blades, while lateral sur-
arches, typically concentrating in the first gill arch [51, faces were smooth. The number of gill rakers decreased
52]. Gill rakers served a dual function: they redirected from the first to the fourth-gill arch in both species, with
Alsafy et al. BMC Zoology (2025) 10:3 Page 7 of 21

sea bream having 14 to 8 lateral and medial gill rakers, In Puffer fish [45], the gill rakers are located on the
respectively, on successive gill arches, and sea bass hav- concave internal side of the gill arches. Grossly, each gill
ing approximately 20, 18, 16, and 12 gill rakers on the first arch has two rows of gill rakers: small lateral gill rakers
through fourth gill arches respectively [4]. (1 mm) and slightly larger medial gill rakers (2 mm). The
Moreover, in Chaca chaca, unlike the gill arches of gill rakers of adjacent gill arches are interdigitated. The
other fish species, there are no gill rakers. This adaptation number of gill rakers on the lateral rows of the 1st, 2nd,
is directly linked to the feeding behavior and dietary pref- and 3rd gill arches is 17, 12, and 11, respectively, while
erences of the species, as first reported by [12]. the number on the medial rows is 17, 15, and 12, respec-
tively (Table 1).
Marine water fish (Table 1) In grey gurnard [15], each gill arch carried two rows
In Siganus luridus [20], the gill arches were L-shaped. of gill rakers: long lateral and short medial gill rakers on
Each gill arch had two different forms of gill rakers with the first gill arch, while the rest of the gill arches showed
asymmetrical arrangements on most parts of the gill short lateral and medial gill rakers. In the striped-red
arches: spine-like gill rakers on the rostral side, which are mullet, all three-gill arches carry short lateral and medial
bifid or trifid spines, and duck toe-shaped rakers on the gill rakers. The gill rakers of adjacent gill arches are inter-
caudal side. The mean number of gill rakers on the 4th- digitated. The average number of gill rakers varies slightly
gill arches is 19, 17, 13, and 10 (Table 1). In Boops boops among gills in grey gurnard and striped-red mullet (Table
[20], the gill arches were semilunar in shape. The first gill 1).
arch carried conical gill rakers with pointed ends on the In Mugil cephalus [64], the lateral row has long and
lateral side, while the gill rakers on the medial side of the numerous gill rakers, while the medial row has shorter
1st gill arch and the medial and lateral sides of the 2nd, and fewer gill rakers (Table 1).
3rd, and 4th-gill arches were short and had spines. The
mean number of gill rakers on each gill arch was 21, 16, Freshwater fish (Table 2)
14, and 13 on the 1st to the 4th gill arches, respectively In Bagrus bayad [3], the first two gill arches had only lat-
(Table 1). eral rakers. The first gill arch carried well-developed long
In Pagrus pagrus [20], the gill arches look like a hook gill rakers, while the gill rakers on the subsequent rows
and carry short gill rakers with fine-needle spinules cov- were short. Most gill rakers were short, an adaptation to
ering the gill rakers’ top, arranged in medial and lateral the carnivorous feeding habits of Bagrus bayad (Table 2).
rows. The gill rakers of the first three gill arches are par- In Tilapia zillii [48], the gill arch carried two rows of
allel, while on the 4th-gill arch, the gill rakers of the lat- small, short, wide-spaced gill rakers: lateral and medial.
eral row correspond to only three small gill rakers on the The gill rakers on each row were nearly the same size
medial row. The mean number of gill rakers is 13, 9, and except for the lateral gill rakers of the first-gill arch, which
9 on the lateral and medial sides of the 1st, 2nd, and 3rd- were longer than the medial ones. Conversely, the medial
gill arches, respectively, while on the 4th-gill arch, the lat- gill rakers of the fourth-gill arch were longer than the lat-
eral gill rakers are 6, and the medial row has 18 gill rakers eral ones. The gill rakers of the adjacent gill arches inter-
(Table 1). digitated with each other. The lateral gill rakers on each
In Sparidae (Sparus aurata and Boops boops) [64], the gill arch were directed dorsolaterally, while the medial
gill arch was bow-shaped with long and more developed ones were directed dorsomedially (Table 2).
gill rakers in the first row and short, less developed gill In Oreochromis niloticus [29], the gaps between the gill
rakers in the second row. The gill rakers in the anterior arches are generally narrow. The gill rakers of the medial
row of the first-gill arch in most species of the fam- row were directed dorsomedially, while those of the lat-
ily Sparidae were short, conical in shape, and elongated eral row were directed dorsolaterally. The gill rakers on
thick strips with slightly pointed ends and triangular the same gill decreased in size towards the epibranchial
bases in Boops boops. part. They appeared relatively short and wide-based with
In European hake [17], the first-gill arch had long lat- tuberous ends. In Chrysichthys auratus [29], the gill rak-
eral gill rakers that appeared triangular with pointed ends ers had two rows; the medial row was directed ventrome-
and carry spines, while the rest of the gill rakers in the dially, while the lateral row was directed ventrolaterally.
row were short with blunt ends and carry spines. The The gill rakers appeared relatively short and broad-based
number of gill rakers on the lateral side of the four gill with segmented tuberous ends (Table 2).
arches is 10, 7, 5, and 3 on the first, second, third, and In Clarias gariepinus [29, 46], the gill rakers were in
fourth-gill arches, respectively, while the number of three rows: medial, intermediate, and lateral. The medial
medial gill rakers is 7, 6, 4, and 2, respectively. The gill and lateral row gill rakers were numerous with long pro-
raker length decreases from the first to the fourth gills cesses, while those of the intermediate row were few and
(Table 1). short. The medial row was found only in the third and
Table 1 Marine water fish studies on varied fish families, feeding types, number and shapes of the gill arches and gill rakers, and the references supported that
Fish Family Feeding type No. gill Gill rakers number Shapes of gill arches Shapes of gill Refer-
Alsafy et al. BMC Zoology

arches 1L 1M 2L 2M 3L 3M 4L 4M rakers ences

Siganus Siganidae Planktivores 4 pairs 19 ± 0.32 17 ± 0.45 13 ± 0.45 10 ± 0.89 Bifid or trifid spines, or duck toe-shaped gill rakers. L shape [20]
luridus
Boops boops Sparidae Omnivorous 4 pairs 21 ± 0.89 16 ± 0.32 14 ± 0.32 13 ± 0.89 1L = long conical gill rakers with sharp lateral sides. Semilunar [20]
1M to 4M had short spiny gill rakers.
(2025) 10:3

Pagrus pagrus Sparidae Carnivorous 4 pairs 13 ± 0.45 9 ± 0.32 6 ± 0.9 18 ± 0.4 Short, had fine-needle spinules covering the gill rakers’ Hook [20]
top.
Sea bass Serranidae Carnivorous 4 pairs 29 26 25 22 17 14 11 9 1L = long saw-like, the rest was short and smooth. Semilunar [4]
Sea Sparidae Carnivorous 4 pairs 11 11 10 10 10 10 8 7 Short, wide-based processes with sharp ends and blunt Bow-like [4]
bream = Spa- surfaces.
rus aurata Short, conical in shape, adapted to carnivorous feeding. [64]
Red Sea Sparidae Carnivorous 4 pairs Short, conical in shape. Semilunar [64]
seabream
(Diplodus
noct)
Dusky Serranidae Carnivorous 4 pairs 28 28 27 27 19 18 17 17 1L = 1ong cone shape, while the reset of gill rakers was Semilunar [19]
grouper cylindrical, short gill rakers.
AC 27 AC 26 AC 17 AC 13 The accessory rakers were like the short gill rakers but
smaller in length.
European Merlucciidae Carnivorous 4 pairs 10 7 7 4 5 4 3 2 1L was triangular with pointed ends carrying spines, ——— [17]
hake with the rest of the gill rakers elevation with blunt ends
carrying spines.
Grey Gurnard Triglidae Carnivorous 4 pairs 16 17 14 14 12 12 11 11 Long lateral and medial short gill rakers on the first gill Crescentic-shape [15]
Fish arch, while the rest gill arches show short lateral and
medial gill rakers in the grey gurnard.
Striped Red Mullidae Omnivorous 3 pairs 11 12 9 10 8 11 - - All three-gill arches were carrying short lateral and Crescentic-shape [15]
Mullet Fish medial gill rakers.
Puffer Fish Tetrodontidae Omnivorous 3 pairs 17 17 12 15 11 12 - - The gill arch has two rows of small lateral and slightly Somewhat cres- [45]
larger medial gill rakers. centic -shape
Mugil Mugilidae Detritus feeder 4 pairs The lateral row had long and great numbers of gill rak- Crescentic [64]
cephalus ers, while there were short and fewer numbers on the -shaped
medial one.
Page 8 of 21
Alsafy et al. BMC Zoology (2025) 10:3 Page 9 of 21

Table 2 Freshwater fish studies on varied fish families, feeding types, number of the gill arches and gill raker, and the references
supported that
Fish Family Feeding type No. of gill Gill rakers characteristics and another characteristic features Refer-
arches ences
Oreochromis Cichlidae Herbivorous 4 Semilunar in shape, the gaps between the gill arches were generally [29]
niloticus narrow. The gill rakers of the medial row were directed dorso-medi-
ally, while those of the lateral row were directed dorso-laterally. The
gill rakers appeared as relatively short and wide-based processes with
tuberous ends.
Chrysichthys Claroteidae Omnivorous 4 The gill arch was semilunar in shape. [29]
auratus The gill rakers had two rows; the medial row was directed ventro-
medially, while the lateral row was directed ventro-laterally. The gill
rakers appeared as relatively short and broad-based processes with
segmented tuberous ends.
Clarias Clariidae Omnivorous 4 and additional branched bulbous dendritic structures originating from the second [29,
gariepinus 5th rudimentary and fourth-gill arches. The gill rakers were presented in three rows: 46]
medial, intermediate, and lateral. The medial and lateral row gill rakers
were numerous with long processes, while those of the intermediate
row were few and short. The medial row was found only in the third
and fourth gills, while the intermediate row was found in the four
main gills,
Bagrus bayad Bagridae family Carnivore 4 Absent medial rows of gill rakers at the first and second gill arches, [3]
the number of gill rakers ranged from 14 to 16. The first gill arch
carried well-developed long gill rackers, while the gill rackers on the
following rows were short. Crescentic outlines with a wide interbran-
chial septum,
Nile tilapia Cichlid Changed to 4 The length of the gill rakers increased from the first to the fourth gill. [29,
Omnivorous The number of the gill rakers was variable in the different gills: they 46]
from herbivo- were 28 in the first gill, decreased to (25) in the second and third gills,
rous with an but increased again to reach a maximum value (38) in the fourth gill.
increase in size
Tilapia Zilli, Cichlid Herbivorous 4 Each gill arch carried two rows of the small, short, wide-spaced gill [48]
redbelly rakers: lateral and medial. The gill rakers on each row were nearly the
tilapia same in size, except the lateral gill rakers of the first-gill arch were
longer. The gill rakers of the adjacent gill arches were interdigitated
with each other. The lateral gill rakers on each gill arch were directed
dorsolaterally, while the medial ones were directed dorsomedially.
Common Cyprinidae Omnivorous 4 The medial edges of four gill arches with zippers have the potential [49]
carp (Cyprinus to interlock. A significant reduction in gill arch length was observed
carpio) from the first to the fourth, with the second and third gill arches
having the greatest number of gill rakers (2nd (25.38 to 29.00) and 3rd
(25.87 to 28.62). The length of gill raker increases from the first to the
fourth gill arch.

fourth gills, while the intermediate row was in all four direction. The gill filaments appeared long in the middle
main gills. The number of gill rakers in the intermediate and shortened toward the extremities [4]. The effective-
row was 20, 23, 19, and 18 in the first, second, third, and ness of fish gills in extracting oxygen from water, as well
fourth gills, respectively (Table 2). as their adaptation to water and immunity, was reported
In common carp, the number of medial row gill rak- by [4, 65].
ers from the first to the fourth gill arch at the right gill
was (25.50 ± 2.45, 27.50 ± 2.78, 27.37 ± 2.61, 21.62 ± 2.00), Scanning electron microscopy of gills
while their lengths were (1.80 ± 0.18 mm, 2.15 ± 0.34 mm, Gill arch
2.38 ± 0.30 mm, and 2.47 ± 0.28 mm) [49]. All surfaces of the gill arch are covered with a mosaic of
irregular polygonal pavement cells with apparent, con-
Gill filaments centrically arranged surface ridges [4]. Microridges,
Each gill arch carried double rows of well-developed resembling fingerprints, covered the exposed surfaces
and compactly arranged gill filaments. Each gill fila- of the epithelial cells. The mucous pores varied in size,
ment row was called a hemibranch, and the two hemi- appearing as narrow, deep, rounded-to-oval holes with
branchs together formed a holobranch. The hemibranchs little or no visible internal structure. Ovoid grooves were
of the four-gill arches were numbered in a lateromedial observed on the dorsal third of the two gill arch surfaces
Alsafy et al. BMC Zoology (2025) 10:3 Page 10 of 21

between the roots of the gill rakers, showing short, sharp, internal structures. There were two types of pores: chlo-
pointed, curved spines protruding through the gill epi- ride and mucus cells with their secretions.
thelium of sea bass (Fig. 3C, D). Taste buds were seen In Tilapia zillii [48], the middle part of the gill rakers
between the gill rakers on the gill arch surface, each dis- did not have any characteristic structures but had two
tinguished by closely packed sensory protrusions towards different elevated structures near the gill filament: an
the surface (Fig. 3C, D). oval-shaped leaf-like structure and a rounded-shaped
In the dusky grouper [19], the upper third of the gill structure. The elevated oval leaf-like structure had a
arch height contained a longitudinal band with many round end carrying two lateral rows of triangular pointed
wavy folds, irregular mosaic patterns, and polygonal epi- spines separated by a median groove, which had a few
thelial cells. The surface of the gill arch between the gill type I taste buds. The elevated, rounded-shaped structure
rakers bases had many small spines and taste buds. On consisted of two round-shaped structures connected to
the dorsal surface of the gill arches, there was a circular each other. The surface of the four-gill arches was cov-
group of spines, ranging from 5–8 spines with medially ered with a mosaic of irregular polygonal pavement cells,
curved ends, located medially to the base of the main with numerous pores of different sizes for chloride and
gill rakers or between them (Fig. 4D, E). Each spine had mucous cells. Type I taste buds were present on the gill
a pointed arrow cape-like structure on its apex, flanked arch.
distally with an annular groove, and the bases of the In three types of fishes from the Nile River (Oreo-
spines had an annular groove. The surface of the gill arch chromis niloticus, Chrysichthys auratus, and Clarias
had a stone-like background texture. gariepinus), the epithelium covering the gill arches had a
The surface ultrastructure of gill arches and gill rakers mosaic of variable dimensions [29]. The exposed surfaces
was derived from studies on fish species having different of the epithelial cells were covered by micro ridges, which
feeding habits: Rhinomugil corsula [66], Gadusia chapra were straight and compactly arranged in Oreochromis
[67], filter feeder Brevoortia tyrannus [68], Hypostomus niloticus but parallel to each other or irregularly inter-
commersonii [69], Prochilodus scorfa [70], Mugil curema, woven to form web-like patterns in Chrysichthys aura-
Mugil liza and Mugil platanus [31], omnivorous Fun- tus and Clarias gariepinus. Mucous cells were scattered
dulus heteroclitus [43], Cyprinus carpio [71], carnivo- among the gill arch epithelial cells, being more numer-
rous Anabas testudineus [66], Notopterous chitala [67], ous in Chrysichthys auratus and Clarias gariepinus than
Eugerres brasilianus [72], Cathorops strigosa [73], car- in Oreochromis niloticus. These mucous cells were often
nivorous catfish, Rita rita [42], argentinian silverside [74], filled with blobs of mucous secretion.
Sakhalin trout [75], snow Trout Schizothorax [76], and In common carp [49], a median crest is demonstrated
Indian major carp Cirrhinus mrigala [77]. on the gill arches between the lateral and medial gill rak-
The surface ultrastructure of the gill arches and the gill ers. The interbranchial septum had mucosal folds that
rakers of an herbivorous fish, the Indian major carp Cir- ran from cranial to caudal and finger-like papillae near
rhinus mrigala, have closely lying short gill rakers and the pharyngeal teeth and replaced mucosal folds. Type II
narrow inter-gill rakers channels on the gill arches, which and III taste papillae were found in the anterior pharynx.
were associated with filtering and retaining food particles
[77]. The surface of the gill arch in the grey gurnard was Gill rakers
characterized by gill rakers with multiple small spines. In sea bream [4], the gill rakers appeared as short, wide-
In the striped-red mullet, the surfaces of the gill arch based processes with sharp ends and blunt surfaces.
appeared smooth, except for a region with many longitu- Many sharp-pointed spines protruded on the gill rak-
dinal micro ridges demarcating the area between the gill ers’ ventral border, while the dorsal border was free from
rakers and the origin of the gill filaments [15]. Addition- spines. In sea bass, the first gill arch had long cylindri-
ally, the striped-red mullet had many taste buds on the cal gill rakers with tapering ends. The medial surface of
smooth surface of the gill arch. these gill rakers carried conical spines of different sizes
In the European hake [17], the surface of the gill arches and directions, while the lateral surface was smooth. The
was wrinkled in some areas. The lateral and medial sur- second type of gill rakers showed a short cylindrical mass
faces of the first gill arch differed from those of the other of spines surrounded by a deep, circular groove. Taste
gill arches. The first-gill arch had a longitudinal line of buds were noticed at the summit of the epithelial protu-
spines presented in groups, forming circular, cuboi- berances, at various elevations between the spines on the
dal, rectangular, oval, and triangular shapes with spaces second type of gill rakers.
between them. In Bagrus bayad [3], the gill arches are In the dusky grouper [19], each gill arch had two main
entirely covered by pavement cells. The pavement cells rows of gill rakers (lateral and medial) and two accessory
had circular and oval openings of variable sizes without rows of gill rakers arranged alternately with the medial
and lateral gill rakers. The two main rows were nearly
Alsafy et al. BMC Zoology (2025) 10:3 Page 11 of 21

similar in length on the second, third, and fourth-gill spines adapted for sorting plankton, similar to observa-
arches, while on the first gill arch, the lateral row had tions by [64]. Additionally, wedge-shaped spines at the
long gill rakers, and the medial row had short gill rak- base of gill rakers act to increase the seizing of slippery
ers. Spines were especially prominent on the long gill and smooth prey [78].
rakers. The lateral gill rakers of the first gill arch had a The gill rakers of Pagrus pagrus were short with fine-
long cone-shaped part, smooth on the lateral surface needle spines covering their tops. The height of the gill
and spined on the medial surface, carrying three types of rakers gradually decreased from the first to the fourth-
spines: long, medium, and short. The small spines on the gill arch. The spines were conical with pointed, curved,
medial aspect of the lateral gill rakers were surrounded or straight ends, confirming the carnivorous nature of
by an annular groove, with taste buds and mucous pores Sparidae fish [53, 64, 83]. Pagrus pagrus had a low num-
arranged in a linear state. The medial row of the first gill ber of gill rakers but a larger number in the last row,
arch and the rows of the following gill arches had cylin- which may increase the seizing of slippery, smooth, and
drical, short gill rakers. The base of the medial gill rak- slimy prey [62].
ers of the first gill arch was circular. In contrast, the gill The arrangement of gill rakers in medial and lateral
rakers’ bases of the medial and lateral gill rakers of the rows in Pagrus pagrus, Boops boops, and Siganus luri-
following gill arches were compressed cylindrical with a dus plays a crucial role in their feeding habits. The equal
dorsal extension on the gill arch. The short gill rakers had number of gill rakers on the medial and lateral rows in
small spines on their bases and bodies, with moderate all gill arches, except in the last gill raker row of Pagrus
and long spines emerging from the epithelial covering. pagrus, is a key factor in their feeding strategy. The high-
The short spines had small openings at their bases with est number of gill rakers on the first gill arch, particu-
a wavy epithelial covering and taste buds’ protrusions, larly in Boops boops, Siganus luridus, and Pagrus pagrus,
while taste buds III were sunken in the epithelium with enhances cross-flow filtering and limits the escape pos-
sensory protrusions. The accessory gill rakers resembled sibilities of small prey [52, 64, 84, 85]. The relationship
the short gill rakers but were smaller and had three types between prey size and the gill raker gap and standard
of spines. The three types of spines were measured, with length is a fascinating correlation that underscores the
the long spine on the apex of the short gill rakers being importance of gill raker characteristics in the feeding
longer than the long spines of the long lateral gill rakers habits of fish species [86]. The herbivorous Siganus luri-
of the first gill arch (Fig. 4D). dus had many gill rakers and narrow spaces between
The gill rakers of Siganus luridus were relatively them. In contrast, the carnivorous Pagrus pagrus had
smooth, ending in spine-like structures with different fewer gill rakers and wider spaces, and the omnivorous
shapes: single spine, bifid, trifid, and quadrate, resem- Boops boops had an intermediate range. This indicates
bling duck toes, which act as filters to catch algae par- that herbivorous species prefer smaller food particles,
ticles. Adapted to vegetarian feeding, Siganus luridus while carnivorous and omnivorous species prefer larger
progresses from feeding on zoo and phytoplankton as food particles.
larvae to finer algae as adults [64, 78]. They primarily Short and tuberous gill rakers in Oreochromis niloticus
consume algae (99.73%), seagrass, and rubble [79]. For were effective filters of food. In Chrysichthys auratus, gill
herbivorous fish, the gill rakers were mainly short, act- rakers were short with a broad base, which strain water
ing as branchial sieves to efficiently filter small food par- to bathe the gills and prevent solid particles from pass-
ticles from the water [77]. The herbivorous black fish’s ing over them. Gill rakers in Clarias gariepinus were
gill rakers direct water toward the oral cavity roof, where long, cylindrical, and arose at acute angles to the gill arch,
the mucous covering traps food particles before being helping to strain food and other materials, thus protect-
ingested [80]. It is suggested that gill rakers perform a ing gill filaments from damage [29].
dual function: changing the direction of the water and fil- The gill rakers and spine distribution on the first gill
tering food particles [80]. arch of the European hake differed from that of the
Boops boops had long gill rakers appearing conical other three gill arches on the lateral and medial surfaces
with pointed ends on the medial and lateral sides of the [17]. In Tilapia zillii [48], scanning electron microscopy
first-gill arch. The following gill arches had long gill rak- showed the gill rakers on both rows were short and small.
ers on their medial sides and short gill rakers on their lat- Each gill raker had a median central axis surrounded by
eral sides. All gill rakers carried various shapes of spines. two lobulated lateral regions. The median central axis
These long and short-spined gill rakers, and the narrow was smooth with a blunted round end, and, under higher
spaces between them, are specialized for different types magnification, its surface was covered with a mosaic
of food particles, as Boops boops are omnivorous [53, of polygonal pavement cells with clear, concentrically
64, 81, 82]. The long gill rakers were semi-conical with arranged surface cell ridges, giving them a fingerprint
tapered ends, and their medial surfaces had needle-like appearance. Oval pores of the chloride cells and a few
Alsafy et al. BMC Zoology (2025) 10:3 Page 12 of 21

small-sized taste buds type I were present. The two lobu- sensory protrusions towards the surface in see bass
lated lateral areas carried numerous pointed spines, taste (Fig. 3C, D). Additionally, the striped-red mullet had
buds, and a few chloride cell pores. many taste buds on the smooth surface of the gill arch
In common carp [49], the lateral and medial gill rak- [31]. In Tilapia zillii [48], a few small-sized type I taste
ers had different numbers of dome-like projections on buds were present on the gill arch. In sea bream [4], the
the side facing the oral cavity. Each gill raker also ends taste buds were noticed at the summit of the epithelial
in a conical tip. The surface epithelium was composed protuberances, at various elevations between the spines
of irregular, polygonal-shaped cells. Microridges were on the second type of gill rakers. In the dusky grouper
found to be arranged differently on cells. Two micror- [19], The small spines on the medial aspect of the lateral
idges were discovered: elongated microridges and short gill rakers were surrounded by an annular groove, with
microvillus-like. type I taste buds, while taste buds III were sunken in the
epithelium with sensory protrusions.
Gill filaments In general, the presence of taste buds on the gill fila-
The gill filaments carried leaf-like lamellae that arose ments has not been reported in any fish species. How-
from both sides. Under higher magnification, the surface ever, a recent study by [13] documented the presence of
of the gill filaments and lamellae was covered by a mosaic taste buds on the gill filaments of the Indian Moth cat-
of irregular, polygonal epithelial cells. These cells exhib- fish, Hara hara, for the first time. They associated this
ited many concentric microplicae, which appeared large adaptation with the food and feeding habits of the fish.
at the bases of the lamellae. Numerous large, rounded to
oval openings of chloride cells with projections, as well as Light microscopy of gills
smaller openings of mucous cells, were observed [4]. Gills consist of gill arches that bear gill lamellae, both pri-
In the dusky grouper [19], each gill arch carried two mary and secondary, supported by hyaline cartilage and
rows of gill filaments on its ventral side. The gill filaments covered by mucous epithelium. Branchial arteries run
were measured at three levels (middle and two extremi- longitudinally through the gill arches, sending an afferent
ties of each gill arch), showing that the gill filaments arteriole to the gill filaments (Figs. 5, 6).
were longest at the middle of the gill arch and shortest at The squamous epithelial cells of the gills were found
the rostral end. The bases of the gill filaments exhibited to have species-specific ridge patterns on their outer
irregular folds with stony-like projections. The primary surfaces [87]. These have previously been misidentified
gill filaments had secondary lamellae that contained the as microvilli or stereocillia. Euryhaline species, such as
pores of mucous and chloride cells. The mucous pores Tilapia zillii, have more dense and well-developed ridges
were larger in diameter than the chloride cell pores. The than stenohaline species, like Sarotherodon niloticus.
secondary lamellae had pavement cells on their surfaces, Freshwater-adapted individuals of Tilapia zillii, Sarother-
with fingerprint-like epithelial coverings and characteris- odon niloticus, Sarotherodon galflaeus, and Tristramella
tic wavy folds. sacra exhibit slightly swollen surface cells that extend
On the surface of the gill filaments of the European over chloride cell openings. During adaptation to sea
hake, longitudinal ridges were present, carrying pores of water, these ridges rise and become denser, while the cell
chloride and mucous cells [17]. surface shrinks, revealing the underlying orifices of the
chloride cells’ apical crypts. The more euryhaline the spe-
Microridges and sensory organs cies, the less chance there is in the ridge pattern of the
All surfaces of the gill arch are covered with micror- cells during the passage from fresh to seawater. This evi-
idges in teleost fish, resembling fingerprints, covered the dence implicates the gill epithelium and the chloride cells
exposed surfaces of the epithelial cells [44]. The gill arch in the process of osmoregulation [87].
has circular microridges can fix the mucus on the surface,
and the strength of cells can be enhanced by the structure Epithelia of the gill arches
of circular microridges that can alleviate the mechani- The structure comprises multiple layers, including super-
cal damage of water flow. Microridges were found to be ficial pavement cells, large chloride cells, goblet cells
arranged differently on cells. Two microridges were dis- secreting mucus, and basal epithelial cells. Pavement cells
covered: elongated microridges and short microvillus- exhibit a cuboidal morphology.
like in common carp [49]. Under light microscopy (Fig. 7), the outer surface of
The surface of the gill arches, gill rakers, and gill fila- pavement cells shows a striated appearance due to micro
ments exhibit various types of taste buds [3, 4, 17, 29– ridges, which may be microvilli or microplicae. Chloride
31]. The gill arch taste buds have a role in food selectivity cells, known as ionocytes, are characterized by their large
[19]. Taste buds were seen between the gill rakers on the size and granular cytoplasm rich in mitochondria (Fig. 8).
gill arch surface, each distinguished by closely packed Mucous cells contain vacuoles, observed in H&E staining
Alsafy et al. BMC Zoology (2025) 10:3 Page 13 of 21

Fig. 5 The paraffin sections of the gills of the silver carp stained by HE stains (A) and PAS stain (B). The gills consist of gill arch, gill filament (primary gill
lamellae, arrowheads) and the secondary gill lamellae(arrowheads). Note arrow heads on figure B pointed out to rodlet cells

Fig. 6 A paraffin section of the silver carp’s gills stained with PAS. The gills
comprise the gill arch, the gill filament (primary gill lamellae), and the sec-
ondary gill lamellae. The cartilage that is heavily stained by PAS supports
the gill filaments

(Fig. 8), and show metachromatic staining properties

with toluidine blue (Figs. 7, 8, 9).
Basal epithelial cells directly contact the basal lamina
(Fig. 8) and serve as progenitors for other cell types. The
lymph space within the gill arch epithelia contains cells
resembling tenocytes, along with immune cells such as Fig. 7 Displays semithin sections of the gill arch of the Ruby-Red-Fin
Shark, also known as the Rainbow Shark (Epalzeorhynchos frenatum),
lymphocytes and granular leukocytes (Fig. 8), rodlet cells which belongs to the Teleostei: Cyprinidae family. The epithelia of the gill
(Fig. 9), and eosinophilic granular cells. arch contained goblet cells, chloride cells, and pavement cells. Goblet cells
Gill arches also feature sensory structures called neu- exhibit a high concentration of metachromatic mucus production. The
romasts, situated on elevated papillae. These structures pavement cells had apical striations, which correspond to microvilli. The
are composed of sensory hair cells and supporting cells, chloride cells possess cytoplasm with granules and mitochondria
including stem cells (Figs. 10, 11).
intermediate cells occupy the space between [89]. Nota-
Filament epithelium bly, the interlamellar area of the gill filament epithelium
The gill filament epithelium resembles gill arches, com- contains numerous large chloride and mucous cells. Both
prising pavement cells, basal epithelial cells, mucous basal and intermediate undifferentiated cells, character-
cells, and chloride cells rich in mitochondria [88]. The ized by a high nucleus-to-cytoplasm ratio, are present in
surface is covered with cuboidal pavement cells, undif- the epithelium of both gill arches and gill filaments. They
ferentiated basal cells contact the basal lamina, and
Alsafy et al. BMC Zoology (2025) 10:3 Page 14 of 21

Fig. 8 Shows a semithin section of the gill arch of the Ruby-Red-Fin Shark, Fig. 10 Illustrates the presence of neuromast on the gill arch of a cat-
also known as the Rainbow Shark (Epalzeorhynchos frenatum), which be- fish, as indicated by the arrowhead. The goblet cells exhibited pale and
longs to the Teleostei: Cyprinidae family. The gill arch epithelia consists of vacuolated cytoplasm, with eosinophilic granular cells in the lymph space
various cellular kinds. Goblet cells exhibit a high concentration of meta- (arrow). The stratum compactum (SC) is located underneath the basal
chromatic mucus production. The pavement cells exhibited apical stria- lamina
tions, which corresponded to microvilli. Telocyte-like cells, indicated by
the arrow, comprise the boundary of the lymph space (L). Lymphocytes,
indicated by double arrowheads, and granular leukocytes, indicated by ar- gills. Lamellae are arranged like beads on a string. Pave-
rowheads, are present in the lymphatic space. Note Basal epithelial cells (b) ment cells form the outer layer of the epithelium and are
characterized as squamous cells. The lamellar epithelium
typically lacks mucous and chloride cells [91].
The structure and function of the branchial chloride
cell in freshwater fishes [92]. Meanwhile, he stated that
the pavement cell is the site of Na+ uptake via channels
that are electrically connected to an apical membrane
vacuolar H+-ATPase (proton pump). Chloride cells play
a critical role in acid-base regulation. Alkalosis increases
the surface area of exposed chloride cells, improving
base equivalent excretion by increasing the rate of Cl−/
HCO3− exchange. In contrast, the chloride cell surface
area is reduced during acidosis due to the expansion of
adjacent pavement cells. This response decreases the
number of functional Cl−/HCO3− exchangers [92].

Gill arch structure

Fig. 9 Displays a semithin section of the gill arch of the Ruby-Red-Fin The gill arch comprises hyaline cartilage in cup-like for-
Shark, scientifically known as Epalzeorhynchos frenatum, a Teleostei: Cy- mations, each consisting of a central cartilaginous core
prinidae family member. The epithelia of the gill arch contained Goblet surrounded by a peripheral matrix. Gill filaments extend
cells that possess a significant abundance of metachromatic mucus pro-
from these gill arches, each containing a cartilaginous bar
duction. Granular rodlet cells are situated within the lymphatic space (L)
covered by an epithelial layer with scattered mucous cells
and sparse chloride cells in the interlamellar epithelium.
serve as precursors for specialized cell types (pavement The extending epithelium covers contralateral lamellae,
cells and mucous-secreting cells) (Figs. 12, 13). while internal lamellae consist primarily of blood spaces
[4].
Lamellar epithelium The gill arches of the European hake are composed of
The lamellar epithelium is thinner than the gill filament hyaline cartilage arranged in cup-like structures with
epithelium and supported by the basal lamina [90]. To a central cartilaginous core and peripheral matrix. Gill
enhance gas exchange, it minimizes blood-to-water dif- filaments extend from these gill arches, which are com-
fusion distances. Pillar cells, specialized endothelial cells posed of a cartilaginous bar with a peripheral matrix
forming blood gaps within the lamellae, are unique to fish and central core. They are covered by an epithelial layer
Alsafy et al. BMC Zoology (2025) 10:3 Page 15 of 21

Fig. 11 Identification of lysosomal activity of rodlet cells (arrowheads) on paraffin section of the gills of sliver carp using Acridine orange

Fig. 12 Displays a semithin section of the gill filament and secondary gill Fig. 13 Displays a semithin section of the gill filament of the Ruby-Red-
lamella of the Ruby-Red-Fin Shark, formally identified as Epalzeorhynchos Fin Shark, scientifically known as Epalzeorhynchos frenatum. This species is
frenatum, a species belonging to the Teleostei: Cyprinidae family. The gill classified under the Teleostei: Cyprinidae family. T. The gill filament com-
arch is upheld by cartilage containing chondrocytes, situated within a la- prises granular rodlet cells (shown by double arrowheads) and transitional
cuna surrounded by a cartilage matrix (asterisk). Telocyte-like cells (shown rodlet cells (indicated by arrowheads), as well as mature rodlet cells (indi-
by the arrow) are present in the epithelia of the gill arch. These cells are cated by a red arrowhead)
generated within the wall of the lymph space (L). Granular rodlet cells,
shown by arrowheads, are situated within the lymphatic space. The sec- perform various tasks, including osmoregulation and
ondary gill lamella comprises pillar cells between the blood capillaries. The transportation [94, 96], Innate immune cells, and prob-
secondary gill lamella is enveloped by pavement cells ably leukocyte-derived and sensory function [97]. These
cells perform a secretory role [98, 99].
containing few mucous cells and scattered chloride cells
in the interlamellar epithelium. Each gill filament bears Neuroepithelial cells
several leaf-like secondary lamellae on both sides, lined The neuroepithelial cells (NECs) of the fish gill fila-
with epithelial pavement cells, some mucous cells, and ment are morphologically and functionally most simi-
pillar cells [17]. lar to the cells of the neuroepithelial bodies in the lungs
Numerous long-gill raker processes from both sides of of air-breathing vertebrates. In teleosts, neuroepithe-
the gill arch appear in Clarias gariepinus [46]. The lateral lial cells are found on the distal half of the filament. In
row gill rakers are present in all gills, including the fifth, trout, these cells are primarily innervated by non-indol-
while medial row gill rakers are found only in the third aminergic nerves that pick up sympathetic neurotoxins.
and fourth gills, longer in the third. Intermediate row gill The gill filament’s proximal half is made up of isolated
rakers are present in all main gills, most developed in the NECs that are also innervated by intrinsic indolaminer-
third and weakest in the first. gic neurons. The NECs exhibit serotonin-like immunore-
Particular cells called rodlet cells have been found in activity in the granular vesicles packed within the basal
freshwater and marine fish [93, 94]. They are displayed soma, as well as processes that surround non-vascular
in several organs. The respiratory organs, the digestive, and vascular smooth muscles in the gill filament. Apical
vaginal, skin, immunological, circulatory, and skeletal processes from neuroepithelial cells occasionally come
systems, the eye, and the abdominal cavity were among into contact with water on the surface of the gill filament
the organs where they were seen [93–95]. Rodlet cells epithelium [100]. NECS are neurotransmitter-containing
Alsafy et al. BMC Zoology (2025) 10:3 Page 16 of 21

chemosensory cells that are diffusely dispersed within Comparative analysis with mammalian mucosa-associated
a thin epithelial layer of the gill filaments and respira- lymphoid tissue (MALT)
tory lamellae of all gill arches. They are innervated by In summary, the structure and function of GALT in fish
afferent fibers from the central nervous system. Thus, share both similarities and differences with MALT in
hypoxic stimulation of gill NECs appears to trigger the mammalian species. Some of the most conserved struc-
production of adaptive cardiorespiratory reflexes that tures that are GALT analogs to those seen in MALT are
help to maintain O(2) uptake in order to meet metabolic secreting mucosa and organized lymphoid tissues. The
demands [101, 102]. immune responses in mammalian MALT and fish GALT
are quite similar, but differences occur in lymphocytes,
Dendritic organ in catfish M cells, and antimicrobial immunoglobulin production.
The dendritic organ is located in the gill arch of catfish. The function of GALT in fish seems to be part of general
It consists of main stems or smaller branches and end mucosal immunity; thus, the main role of this tissue is to
bulbs comprising elastic cartilage, a vascular layer of con- fight against the pathogens that mainly invade the host
nective tissue, and an epithelium containing intraepi- mucosal surfaces. Stimulation of GALT by vaccination
thelial mucous glands. Blood capillaries originate from has been shown to reduce disease in several common
the vascular layer, penetrate the epithelium, and dilate farmed fish species; thus, understanding of GALT abili-
towards the surface to form respiratory papillae [47, ties is being gradually used for further strategic develop-
103]. The dendritic or arborescent organs that project ment in fish aquaculture at the present time. The cells
into the suprabranchial chamber and are directly derived in the GALT provide surveillance against a broad array
from the second and fourth gill arches [104]. All of the of intruding pathogens. The cells may be lymphocytes,
epithelia in the gill fans, suprabranchial chamber mem- polymorphonuclear cells, monocytes/macrophages,
brane, and labyrinthine organs contain a series of respira- and myeloid cells. These cells are in the lymphoid pop-
tory islets with transecting capillary flow, indicating that ulation in the gill filaments and interbranchial space.
they are derived from the gill filament epithelium. The Mechanisms of immune defense such as pathogen trap-
intraepithelial capillaries extend to the surface, forming ping, inflammation, antigen processing and presentation,
respiratory transverse blood capillaries (papillae) [47]. lymphocyte activation, local immunoglobulin produc-
Based on structural and immunohistochemical con- tion, and NK cell-like activity are also mainly involved in
firmation, it appears that, in addition to the gills, which fighting against a wide variety of pathogens in mamma-
are the primary site for O2 chemoreception in fishes, the lian MALT as well as fish GALT. The types of pathogens
accessory respiratory organs (ARO) in the air-breath- encountered by the MALT in any vertebrate species are
ing catfish (C. gariepinus) act as a functional system for shaped by the environmental factors and the locations in
O2 sensing based on the presence of NEC. NECs have the bodies of the vertebrates [112–114].
also been found in the specialized respiratory epithe-
lia of some catfish species’ accessory respiratory organs Gill-associated lymphoid tissue (GALT)
(AROs), including the gills and skin [105, 106]. Gill-associated lymphoid tissue (GALT) is a multicom-
ponent system distributed throughout the gills of fish. It
Immune system of the gill consists of the pharyngeal region, including tubular ele-
The fish gill’s immune functions are closely linked to gill- ments of the kidney, and a pair of discrete anterior and
associated lymphoid tissue (GIALT) and T lymphocytes. posterior caudal regions. The GALT complex is described
GIALT and T lymphocytes can produce immune mol- as the primary interface between the host and the sur-
ecules that aid gill immunity and protect against patho- rounding aquatic environment. This review encompasses
genic bacteria on mucosal surfaces [107]. In Atlantic the regulation of GALT physiology and the implications
salmon (Salmo salar) [108], the interbranchial lymphoid of GALT in aquaculture research. GALT consists of a
tissue (ILT) was recognized. Consequent studies in Atlan- variety of tubular tissue compartments, including granu-
tic salmon and rainbow trout (Oncorhynchus mykiss) lar and glandular sections, and has a number of defined
demonstrated the presence of T lymphocytes implanted structures that proceed to more distal regions of the gill
in a meshwork of reticulated epithelial cells in ILT. Only a in a reticulated pattern. GALT is composed of MALT ele-
few immunoglobulin (Ig) M+ lymphocytes were noticed. ments, such as macrophages, reticular cells, and plasma
Still, the structure appears rich in cells expressing major cells, arranged within a reticle of crescentic myoepithelial
histocompatibility complex (MHC) class II+ cells and IgT cells. Within the reticle is a labyrinth of capillaries that
transcripts [109, 110]. The ILT must be deliberated as a extends across the gill filaments and is filled with blood in
portion of the gill-associated lymphoid tissue (GIALT), fixed specimens. Two types of benchtop laboratory assays
which is defined as one of the four main mucosal immune have been developed using isolated and enriched gill leu-
compartments found in bony fish [111]. kocytes from rainbow trout as the biological source for
Alsafy et al. BMC Zoology (2025) 10:3 Page 17 of 21

investigating the in vitro production of fish immunoglob- infections in a contagious manner. Therefore, enhance-
ulin from GALT-associated plasma cells. Reports of the ment of gill mucosal immunity in fish is the most direct
presence of toluidine blue-positive cells within gill tissue way to protect against resistance to mucosal pathogens
in fish of several species have suggested that there may be [44, 112, 119–122].
a link between the presence of GALT and the gill mor-
phology of teleosts. Vascular anatomy of the fish gill
Gill-associated lymphoid tissue (GALT) is an important Three vascular networks can be found within the gill
primary inductive site of the branchial immune response filament. The arterioarterial (respiratory) pathway com-
and has unique immunological and anatomical features, prises lamellae, afferent and efferent segments of the
which are likely an adaptation to aquatic life. GALT is the branchial and filamental arteries, and lamellar arterioles.
major production area of gill immunoglobulin T (IgT) in The gill filament has two post-lamellar pathways: inter-
the gill repopulation and is also associated with cell sur- lamellar and nutrient. The interlamellar system is a vast
face markers CD3 or CD8, lymphocyte maturation, and ladder-like network of thin-walled, highly distensible ves-
differentiation that occurs in the gill. GALT contains high sels that runs between and parallel to the gill filament’s
proportions of CD3+ cells among nonadherent leuko- lamellae and around its afferent and efferent borders.
cytes affected by stress application in the mucosal vac- The medial wall of the efferent filamental artery contains
cination site. In the GALT model system, gill fragments short, narrow-bore feeder vessels that supply interlamel-
transplanted from donor fish were transplanted, leading lar vessels [123–126].
to plasma cell, CD4+, and secretory IgT plus mucus epi-
thelial cell infiltration into the recipient’s gill tissue and Conclusion
mucus, which is not specific and will not proceed in a This review comprehensively examines the gill morphol-
time-dependent manner [115–118]. ogy across various marine and freshwater fish species,
highlighting the intricate adaptations to their respective
Function of the gill-associated lymphoid tissue environments and feeding habits. Morphological inves-
Gill, as the principal respiratory organ of fish, has been tigations reveal that the organization of fish gills varies
historically linked to respiration and ion regulation. If widely. The studies reviewed span diverse fish families,
you are an immunologist like me, you must wonder: offering detailed insights into the structural variations
what about the immune functions of the gill? Indeed, gill of gill arches, gill filaments, and gill rakers. We observed
mucosa is rich in immune cells that function not only in significant differences in the arrangement, size, and
immune surveillance but also in maintaining mucosal number of gill rakers between marine and freshwater
homeostasis and protecting fish against pathogens and fishes, reflecting their living and feeding habits. Her-
environmental stressors. Its role as an external defense bivorous fish like Siganus luridus exhibit short gill rak-
barrier is gaining much attention. This section summa- ers designed to filter algae efficiently. At the same time,
rizes the known important immune functions of gill- predatory species such as Pagrus pagrus possess fewer
associated lymphoid tissue and discusses its potential but longer gill rakers with specialized spines for captur-
role in mucosal defense against pathogens. The location ing slippery prey. Omnivorous species like Boops boops
of the gill allows for continuous antigenic sampling that display intermediate gill raker structures allowing ver-
is conducted primarily by intraepithelial lymphocytes. satile feeding strategies. This indicates that herbivorous
The teleost gill contains numerous lymphocyte enrich- species prefer small food particles, whereas carnivorous
ment or induction sites that can effectively guard the gill and omnivorous species tend to consume larger food
against pathogenic invasions. It generates a variety of particles. Microscopic and ultrastructural observations
immune functions, including picking up or recognizing reveal that gill arches are supported by hyaline carti-
pathogens and activating recruited innate leukocytes. In lage and covered by a mucous epithelium rich in various
the adaptive immune system, it regulates the activation, cell types, including pavement cells, chloride cells, and
maturation, and proliferation of gill leukocytes by releas- mucous cells. These cells play vital roles in maintaining
ing cytokines mainly produced by T cells in response to ion balance, mucus secretion, and forming the protective
pathogen stimulation. It also plays a role in transport- and functional layers of the gills. SEM studies underscore
ing antigens to peripheral lymphoid tissue for immune the presence of complex microstructures such as micro
memory induction. Activation of mucosal immunity in ridges and microplicae on epithelial cells, enhancing the
gills is an important factor in fish mucosal defense and surface area for gas exchange and mucus secretion. The
contributes to the overall health and mucosal homeosta- presence of mucous taste buds and immune cells on gill
sis of fish. Fish exposed to live vaccines mainly generate rakers indicates significant surface features contributing
memory T and Ig+ B cells in the gill, so that gill mucosal to the multifunctionality of gill structures. The dendritic
immunity may also strongly protect fish from pathogenic organ in catfish, with its unique structure comprising
Alsafy et al. BMC Zoology (2025) 10:3 Page 18 of 21

Received: 15 September 2024 / Accepted: 20 January 2025

elastic cartilage, vascular connective tissue, and mucous
glands, exemplifies specialized adaptations in freshwater
species for enhanced respiratory efficiency. The diverse
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