Int. J. Aquat. Biol. (2014) 2(3): 147-154 

E-ISSN: 2322-5270; P-ISSN: 2383-0956

Journal homepage: www.ij-aquaticbiology.com 

© 2014 Iranian Society of Ichthyology 

Original Article 

Structural characterization of the olfactory epithelium of freshwater olive barb, Puntius sarana 
(Hamilton, 1822) 

 
Saroj Kumar Ghosh, Padmanabha Chakrabarti*1 

 
Fisheries Laboratory, Department of Zoology, The University of Burdwan, Burdwan-713 104, West Bengal, India. 

 

 

 

 

 

Article history: 
Received 30 May 2014 

Accepted 17 June 2014 

Available online 2 5 June 2014 

 
Keywords:  
Olfactory epithelium 

Function 

Histoarchitecture 

Surface architecture 

Abstract: The cellular organization of the olfactory epithelium of Puntius sarana (Hamilton, 1822) 
was explored by means of optical and scanning electron microscopy. The oval shaped olfactory rosette 

was composed of 26-28 primary lamellae distributed from both sides of the central raphe. The sensory 

epithelium confined chiefly on the linguiform processes of the lamella and rest of the portion consisted 

of non-sensory epithelium. The sensory epithelium was embossed with morphologically distinct three 

types of sensory cells: ciliated, rod and microvillus receptor cells. The non-sensory epithelium was 

made up of mainly stratified epithelial cells and mucous cells. Different cells lining the olfactory 

epithelium were discussed in relation to mode of life and living of the fish concerned. 
 

Introduction 

The olfactory organs are of special attention in fish 

biology because they are basically chemoreceptors, 

efficient to ascertain water soluble compounds and 

afford the fish with information about the nearby 

environment. In fish, water with dissolved chemicals 

flow from anterior to posterior nostril over the 

olfactory organ, covered with olfactosensory 

epithelium which performs an important role in 

olfaction (Hara, 1971). Olfactory information is 

extremely important in the lives of fishes including 

feeding, prey detection, predator avoidance, species 

and sex recognition, and migration (Camacho et al., 

2010). Teleostean olfactory organs exhibit a 

respectable diversity, reflecting different 

developmental strategies and ecological habitats 

(Zeiske et al., 1992). The microstructure and 

function of olfactory organ in several fishes have 

been widely described by several workers (Mandal 

et al., 2005; Sawad et al., 2006; Bhute and Baile, 

2007; Kumari, 2008; Ma and Wang, 2010; 

Chakrabarti and Ghosh, 2011; Ghosh and 

                                                           
* Corresponding author: Padmanabha Chakrabarti 

E-mail address: dr.pchakrabarti@yahoo.in  

Chakrabarti, 2012; 2013; 2014). However, no 

attempt has been made to correlate the functional 

significance of various cells lining the olfactory 

epithelium in relation to the species feeding 

adaptations.  

Puntius sarana (Cypriniformes, Cyprinidae) is 

freshwater, omnivorous, bottom dwelling teleost and 

distributed throughout the Indian subcontinent 

(Chetia Borah, 2012). Its food primarily consists of 

algae, unicellular and single celled animals, larvae of 

aquatic insects and sands (Shafi and Quddus, 2001). 

The aim of the present work is to examine the 

distribution of different cell types and their 

functional aspects in the olfactory epithelium of 

important food fish, P. sarana by histological and 

ultrastructural examination. 

 

Materials and methods 

Living mature specimens of P. sarana (14 to 16 cm 

in total length) were obtained from the local 

freshwater body of Burdwan, West Bengal, India. 

The specimens were anaesthetized with tricane 



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Ghosh and Chakrabarti/ structure of olfactory epithelium in Puntius sarana 

methone-sulphonate (MS 222; Sigma Chemical Co.) 

solution (100 mg/L) and sacrificed following the 

guidelines given by the Institutional Ethical 

Committee. The olfactory organs were carefully 

dissected out from the floor of the nasal cavity under 

a stereo microscope and immediately processed for 

the histological and scanning electron microscopic 

studies. 

Histological study: The olfactory tissues were fixed 
in aqueous Bouin’s fluid for 16-18 hour. After 

fixation the tissues were washed repeatedly in 70% 

ethanol and dehydrated properly through ascending 

series of ethanol. Then they were cleaned with 

xylene and embedded in paraffin wax of 56-58°C 

under a thermostat vacuum paraffin-embedding bath 

for a period of 1 hour and 30 minute.  Serial sections 

were cut at 4 µm thick using a rotary microtome 

(Weswox). After routine histological procedure 

deparaffinized sections were stained with Delafield’s 

Haematoxylin-Eosin (HE) and Mallory’s triple (MT) 

stain. The prepared histological sections were 

examined and photographed by Olympus-Tokyo 

PM-6 compound microscope.  

Scanning electron microscopic (SEM) study: The 
olfactory rosettes were perfused in vivo with 2.5% 

glutaraldehyde solution in 0.1 M Na-cacodylate 

buffer (pH 7.4) for 15 minute. The entire olfactory 

rosettes were dissected out and rinsed with 

heparinised saline (heparin sodium salt 10,000 IU 

dissolved in 0.67% NaCl solution) and 1% Tween 40 

mixture to remove the adhering mucus. After rinsing 

in 0.1 M Na-cacodylate buffer, the tissues were again 

fixed with 2.5% glutaraldehyde in 0.1 M Na-

cacodylate buffer (pH 7.4) for 24 hour at 4°C. After 

primary fixation the tissues were rinsed in the same 

buffer and post-fixed in 1% osmium tetroxide in 0.1 

M Na-cacodylate buffer (pH 7.4) for 2 hour. The 

tissues were washed thoroughly in the same buffer 

and dehydrated through graded series of ethanol 

followed by acetone and isoamyl acetate. The 

samples were dried with critical point drier (Hitachi 

8CP2), mounted on metal stubs, coated with gold 

palladium by vacuum gold coater. The tissues were 

examined under a Hitachi S-530 scanning electron 

microscope. 

 

Results 

According to SEM observations, the olfactory 

rosette of P. sarana is nearly oval in outline with a 

convex ventral and a concave dorsal appearance. The 

olfactory rosette consists of a series of 26 to 28 more 

or less flattened lamellae, distributed in a feather like 

style on both sides of the central raphe and occupies 

the entire cavity of the olfactory chamber (Fig. 1). 

The outer edges of the lamellae are adhered to the 

olfactory chamber, while the inner edges are 

attached to the median raphe. The lamellae are 

capacious in the middle portion of the rosette and 

diminish in sizes towards the preceding and 

succeeding ends. The middle dorsal part of the 

lamella is characterized by free well-developed 

linguiform process, grows from the distal edge 

(Fig. 1). The sensory epithelium is restricted mainly 

on the linguiform processes of the lamella, while rest 

of the portion is consisted of non-sensory epithelium. 

Histologically, the olfactory lamellae are radiated 

from raphe and composed of two layers of thick 

olfactory epithelium separated from the narrow 

central lamellar space, central core by a well-

developed basement membrane (Fig. 2). The central 

core is packed with connective tissue fibers, nerve 

fibers and blood vessels (Figs. 3, 4, 8). The sensory 

Figure 1. Photomicrographs of the olfactory epithelium of 

P. sarana. Oval shaped olfactory rosette exhibits olfactory 
lamellae (OL) radiating from central median raphe (R). Arrows 

indicate linguiform processes of OL (SEM × 50). 



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Int. J. Aquat.Biol. (2014) 2(3): 147-154 

epithelium is typified with mainly receptor cells 

having long dendrite reaching up to the free 

epithelial surface and few rod cells and microvillous 

cells (Fig. 3). Rod cells are ovoid in appearance, 

extend as a spike like texture and characterized with 

basally situated deeply stained nuclei. Microvillous 

cells are situated in more superficial layer of the 

epithelium, possess a round nucleus and without cilia 

(Fig. 3). Receptor cells are the sensory elements of 

the olfactory epithelium and differentiated into 

primary and secondary receptor cells. The primary 

receptor cells are highly basophilic with almost 

rounded and deeply stained nuclei in the distal 

portion of the cell. The dendrite process of each 

receptor cell runs to the superficies of the lamellae as 

a spare cylindrical process (Fig. 4). The secondary 

receptor cells are distributed below the primary 

receptor cells and marked by their oval and 

elongated nuclei. The axonal ends of primary 

receptor cells synapse with the dendrite tips of 

secondary receptor cells (Fig. 4). Under SEM 

observations, the linguiform process of olfactory 

lamella has a spongy structure due to the appearance 

of compact ciliated receptor cells (Fig. 5). The 

Figure 2. Histological sections of the olfactory epithelium of 

P. sarana. Olfactory lamellae (OL) based on raphe (R) shows 
olfactory epithelium (OEP) separated from central core (CC) by 

basement membrane (MT × 100). 

Figure 3. Histological sections of the olfactory epithelium of 

P. sarana. Sensory olfactory epithelium (OEP) lined with receptor 
cells (RC) (solid arrows), rod cells (arrow heads) and 

microvillous cells (broken arrows). Note the presence of blood 

vessels (BV) in CC (HE × 400). 

Figure 5. Photomicrographs of the olfactory epithelium of 

P. sarana. Sensory epithelium exhibits bunch of ciliated receptor 
cells (RC) (SEM × 3500). 

Figure 4. Higher magnification of sensory olfactory epithelium 

(OEP) of P. sarana shows the arrangement of primary receptor 
cells (solid arrows) and secondary receptor cells (broken arrows). 

Note presence of blood vessels (BV) in the central core (CC) (MT 

× 1000). 



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Ghosh and Chakrabarti/ structure of olfactory epithelium in Puntius sarana 

receptor cells are distinguished by the distal free end 

of the dendrites and categorized into ciliated, rod and 

microvillous receptor cells. Ciliated receptor cells 

are furnished by tuft of long cilia, rise to the 

epithelial surface and dominant over both rod cells 

and microvillous cells (Fig. 6). The rod cells are few 

in number and their apical end protrude as a simple 

rod like structure, distributed randomly in the 

epithelial surface. The microvillous cells are 

embossed with tuft of microvilli and dipped into the 

thickness of the ciliated receptor cells (Fig. 6). The 

transitional zone of sensory and non-sensory 

Figure 6. Photomicrographs of the olfactory epithelium of 

P. sarana. Sensory epithelium displays dendrite processes of 
receptor cells (RC), rod cells (solid arrows) and microvillous cells 

(arrow heads) (SEM × 4000). 

Figure 7. Photomicrographs of the olfactory epithelium of 

P. sarana. Transitional zone of sensory and non-sensory 
epithelium shows ciliated receptor cells (RC), microvillous cells 

(broken arrows), stratified epithelial cells (SEC) with complex 

microridges and mucous cells (solid arrows). Note mucin 

droplets over the SEC (SEM × 4500). 

Figure 9. Photomicrographs of the olfactory epithelium of 

P. sarana. Surface epithelium of raphe characterized with 
stratified epithelial cells (SEC) having labyrinth pattern 

microridges. Note the presence of opening of mucous cells (solid 

arrows) in between SEC and mucin mass (arrow heads) over the 

SEC (SEM × 4000). 

Figure 8. Histological sections of the olfactory epithelium of 

P. sarana. Non-sensory olfactory epithelium (OEP) shows 
mucous cells (solid arrows) and stratified epithelial cells (arrow 

heads). Note presence of blood vessels (BV) in the central core 

(CC) which is detached from OEP by basement membrane (BM) 

(HE × 400). 



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Int. J. Aquat.Biol. (2014) 2(3): 147-154 

epithelium is provided with stratified epithelial cells 

intercalated with the opening of mucous cells leaving 

microvillous cells adjacent to the ciliated receptor 

cells (Fig. 7). Histologically, the non-sensory 

epithelium is lined with chiefly stratified epithelial 

cells with prominent basophilic nuclei and mucous 

cells (Fig. 8). Under SEM study, the non-sensory 

epithelium and raphe is represented by tightly 

packed stratified epithelial cells having labyrinth 

pattern microridges and opening of mucous cells. 

Secreted mucin masses are placed over the 

microridges of stratified epithelial cells (Fig. 9).  

 

Discussion    

Olfactory mucosa comprising the sensory neurons is 

typically placed on the floor of the olfactory 

chamber, which is frequently folded, forming 

olfactory lamellae (Hara, 1975). The feeding habits 

of fishes are reflected on the structure and cellular 

organization of the olfactory organ (Hara, 1994). The 

oval shaped olfactory rosette of P. sarana with two 

rows of olfactory lamellae present on both side of the 

median raphe belongs to Burne’s  (1909) rosette 

column I or Bateson’s (1889) rosette type III. 

Teichmann (1954) classified this oval type of 

olfactory organ under the category of eye-nose fish, 

which means that this kind of fish possess similarly 

developed optic and olfactory senses. The olfactory 

organ of P. sarana holds 26-28 lamellae arranged on 

both left and right side of the rosette, adapted 

maximum to the space availability and is similar to 

other cyprinid olfactory organ can be placed to the 

type VI by Yamamoto and Ueda (1979). The 

multilamellar arrangement increases the sensitivity 

and efficiency of the olfactory organ (Zeiske et al., 

1976). The distribution of sensory and non-sensory 

areas in the olfactory epithelium is variable among 

teleosts (Yamamoto, 1982). In P. sarana the sensory 
epithelium is bounded in the free linguiform 

processes, while the rest portion of lamellae is 

comprised of non-sensory epithelium. This 

disposition may be due to the fact that the linguiform 

process with sensory cells faces the flow of incoming 

water current and the receptor cells interact with the 

water soluble chemicals during olfaction. Similar 

findings were also reported by Ojha and Kapoor 

(1973) in the olfactory apparatus of Labeo rohita.  
The most interesting characteristics of the olfactory 

epithelium of P. sarana is the histological existence 
of secondary receptor cells in addition to primary 

receptor cells and the presence of synaptic 

connection between these two kinds of receptor 

cells. This is in conformity with the findings of Goel 

(1978) in the olfactory epithelium of Notopterus 
notopterus. Buck and Axel (1991) reported that 
olfaction commences at the apical tip of the receptor 

cells. In the present observation, the sensory 

epithelium chiefly consists of three morphologically 

distinct types of receptor cells: ciliated, rod and 

microvillous cells; intermingled in different 

proportions. The ciliated receptor cells correspond to 

the type I cells of Yamamoto and Ueda (1978), the 

rod cells similar to the type IV cells of Ichikawa and 

Ueda (1977), whereas the microvillous cells are 

harmonized with the type II cells of Muller and Marc 

(1984). The present study reveals that the ciliated 

receptor cells dominating over the rod cells and 

microvillous cells. The cilia are the site of 

transduction process, stimulated by odour bearing 

substances and also enable the fish to detect food. 

Sparsely distributed rod cells in the olfactory 

epithelium of P. sarana are considered as a sensory 

receptor cells. Yamamoto (1982) opined the rod cells 

as a sign of aging of ciliated receptor cells and 

terminate with single cilium at the free surface of the 

epithelium. Reite and Evensen (2006) believed that 

these are non-specific immune cells, rolled up in 

immunity as their number proliferates during 

parasitic infection. However, on the basis of 

experimental work and developmental studies, 

Zielinski and Hara (1988) and Moran et al. (1992) 

have established that the rod shaped process of these 

cells represent the dendritic apical processes of 

olfactory receptor cells. In P. sarana the microvillous 
receptor cells probably form a different olfactory 

transduction mechanism for pheromones in the 

regulation of reproductive activities. This is in 

compliance with the findings of Biju et al. (2003) in 



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Ghosh and Chakrabarti/ structure of olfactory epithelium in Puntius sarana 

the olfactory epithelium of Cirrhinus mrigala. Bhute 

and Baile (2007) also advocated that the 

microvillous receptor neurons perceive and process 

signals of pheromone, which is an important step of 

breeding in Labeo rohita. On contrary Bakhtin 

(1977) and Bannister (1965) considered that 

microvillous cells in the olfactory surface of Squalus 

acanthias and teleostean fishes are predecessors of 

ciliated receptor cells. 

In the transitional zone of sensory and non-sensory 

epithelium few ciliated receptor cells in between 

stratified epithelial cells are responsible for better 

monitoring of the water quality even up to this zone. 

In P. sarana, the non-sensory cells lining the 

olfactory epithelium may help in mechanical assist 

of sensory cells. Furthermore, the non-sensory 

epithelium and the raphe consist of stratified 

epithelial cells provided with labyrinth pattern 

microridges on their apical surface. Such 

microridges may enhance the epithelial surface and 

serve as holding the mucus film over the epithelial 

membrane and protect the epithelium from 

mechanical abrasion. The mucin secreted by mucous 

cells over the lamellae probably forms a suitable 

medium for diffusion of odorants. The mucus layer 

may also help in ion trap, which obstructs the 

penetration of salts and heavy metals to underlying 

organs (Banerjee, 1993). In addition the mucin 

secreted from the mucous cells of raphe probably 

helps the smooth flow of water through the olfactory 

chamber by binding microscopic debris which is 

ejected through the posterior nostril. This is in 

confirmed with the findings of Bandyopaghyay and 

Datta (1998) in the olfactory organ of 

Heteropneustes fossilis. 

 

Acknowledgement 

The authors are grateful to Dr. A. Basu, Head of the 

Department of Zoology, The University of Burdwan 

for providing laboratory facilities and thankful to Dr. 

S. Chakraborty, The Scientist-in-charge of the 

University Science Instrumentation Centre, The 

University of Burdwan, for his technical support in 

SEM analysis. 

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