Bivalve filter feeding, form, and function

Abstract
There are more than 15000 species of bivalves. Each has its functional morphology that
they have evolved to survive according to its habits and habitat. Bivalves are important
in maintaining the air quality in their environments. In this study, we dissected and
analyze five specimens of five species of bivalves, a rock mussel (Perna viridis), a rock
oyster (Succostrea sp.), a brackish water clam (Geliona expansa), a mud-burrowing
razor clam (Ensis sp.), and an epibenthic scallop (Pectin sp.). The adaptions of shell,
siphons, foot, hinge, adductor muscles, mantle, and organs involved in filter-feeding
were looked at.

Introduction
Bivalves are a group of marine and freshwater mollusks characterized by a shell divided
from front to back into left and right valves connected by two hinged parts known as
adductor muscles. Bivalves include more than 15,000 species of clams, oysters,
mussels, scallops, etc.

Filter feeding is procuring suspended microscopic food particles by randomly straining

them from the water. Filter feeders pump ambient water through a filter that will retain
suspended matter. Most bivalves filter feed using a specialized gill used for breathing
and feeding known as ctenidia. Bivalves filter phytoplankton, bacteria, and other
particles during filter feeding which maintains water quality by transforming filtered
organic matter into tissue or faecal pellet (Petersen et al., 2019). Studying the internal
anatomy of the gills and labial palps will better understand how bivalves filter feed.

Bivalves are found in many aquatic habits throughout the world. Some bivalves burrow
in sediments, others lie on the surface of the seabed or the surface of rocks and hard
surfaces. There are even bivalves that are capable of swimming. These different
habitats and behaviours resulted in a large diversity of bivalves. By investigating the
functional morphology of bivalves, we will be able to show how it relates to their habitat
and behaviour.
The objective of this study is to describe the general morphology of the organs that
bivalves use for filter feeding and the relationship of the functional morphology of
bivalves to habitat and habit. In this study, to get a better understanding of how bivalves
filter feeds a sample from each species, Perna viridis, and Geloina expansa will be
dissected the internal structure of two bivalves. To understand the functional
morphology of bivalves relating to their respective habitats and habits the 5 species of
bivalves will be compared. These 5 species of bivalves chosen are a rock mussel
(Perna viridis), a rock oyster (Succostrea sp.), a brackish water clam (Geliona expansa),
a mud-burrowing razor clam (Ensis sp.), and an epibenthic scallop (Pectin sp.). These
bivalves were chosen because of their different habitats and behaviors

Methods
The specimens for each species were provided by the lab. The Perna viridis specimen
was opened by slicing along the slit between the valves, after doing so the two adductor
muscles were cut at the left valve to separate the two lateral valves. The valves were
then put into a water basin to float the organs. The specimen was determined to be a
make as the gonad was pale white in coloration. Photographs of the left valve were
taken to show where the adductor muscles connected to the valve and the external
features of the shell. Photographs of the right valve are taken to show the arrangement
of internal organs, in one photograph the gonad was lifted to make the gill more visible
(Figure 1).
Fig. 1 Lateral view of Perna viridis. (A) External view of the right valve. (B) Internal view
of the right valve. (C) Internal anatomy of Perna viridis. (D) Internal anatomy of Perna
viridis with gonad lifted.

The Geloina expansa was opened by cutting along the slit between the valves, then the
two adductor muscles were cut at the left valve to separate the two valves. The valves
were then put into a water basin to float the organs. Photographs of the left valve were
taken to show where the adductor muscles connected to the valve and the external
features of the shell. Photographs of the right valve are taken to show the arrangement
of internal organs (Figure 2).
Fig. 2 Geloina expansa. (A) External view of the right valve. (B) Internal view of the right
valve. (C) Internal anatomy of Geloina expansa.

The specimen samples of Succostrea sp., Solen or Ensis sp., and Pectin sp. were all
provided by the lab with the valves separated and the internal organs removed except
Succostrea sp which was submerged in water in a basin. Photographs of the external
features of the shells for each species were taken and so was the right valve of
Succostrea sp. with the internal organs still attached and not attached (Figure 3, 4 & 5).
Fig. 3 Lateral view of Saccostrea. (A) External view of the right valve. (B) Internal view
of the right valve. (C) Internal anatomy of Saccostrea.

Fig. 4 External lateral view of Ensis.

Fig. 5 External lateral view of Pectin.

Results
The valves of Perna viridis are convex, elongated, triangularly ovate in shape, and end
in a ventrally pointing hinge at the umbos terminal (Figure. 1A). The left valve
possesses two small teeth whereas the right valve possesses a single large tooth that
interlocks with each other. Valves have irregularly spaced concentric ridges and on the
left valve, the mantle is fused. Large kidney-shaped adductor muscles are found only
on the posterior side of the shell shown by the adductor muscle scar on the interior of
the right valve (Figure. 1B). The byssus apparatus is located at the base of the foot at
the posterior end, the byssus threads are thick and long (Figure. 1C). The two siphons
located at the posterior end are small in size and are inhalant fringed. The mantle is
fused. The layered ciliated gills of Perna viridis run along the ventral side from the
posterior end to the anterior end. Large labial palps are located at the end of the gills
(Figure. 1D). The foot is small, flat and tongue shaped.

Geloina expansa have bilaterally symmetrical thick valves that are trigonal and rounded
in shape (Figure. 2A). The valves become narrow at the dorsal end where the hinge is
located. The shells are interlocked by articulating teeth at the hinge. On the left valve,
the mantle is fused. Concentric ridges radiate from the umbo terminal to the posterior
ventral margin. Figure. 2B shows that the adductor muscles are located on opposing
sides of the valves, the anterior and posterior sides. The food is large, wide, and long.
At the posterior end, there are two small tubular inhalant fringed siphons. The siphons
are relatively smaller than the average size. Large, layered ciliated gills are hidden
underneath the mantle skirt that runs from the posterior to the anterior (Figure. 2C).
Labial palps are located at the anterior of the gill and are long and narrow.
Saccostrea have laterally asymmetrical valves. The right valve is flat and shorter while
the left valve is rounded and longer, both are irregularly shaped giving it a ‘rocky’
appearance (Figure. 3A). The mantle is fused to the left valve. The hinge is located at
the anterior end and is interlocked by hinged teeth, concentric ridges fan out from the
hinge. The large abductor muscle is located in the middle of the specimen, a valve
(Figure. 3B). Foot and siphons are absent in the specimen. Layers of gills run along the
ventral anterior margins of the valve (Figure. 3C).

Ensis have smooth, elongated, convex, thin, and laterally symmetrical valves, with
concentric ridges radiating from the anterior end to the posterior end (Figure. 4). The
valve is located at the anterior end, it is interlocked by three hinge teeth on the left valve
and two hinge teeth on the right valve. The mantle is fused to the shell. The adductor
muscle is located at the posterior end. The foot of the specimen is large in size. The
siphons are very long and are found on the posterior side.

Pectin has thin, symmetrical fan-shaped valves, though the left valve is more concave
than the right (Figure. 5). There are two wings at the dorsal end, the anterior wing is
always longer than the posterior wing. The hinge is located at the dorsal end and
interlocked together by a hinge ligament called resilium. Radial ridges extend outward
from the umbo to the curved ventral edge of the shell. Its mantle is fused to the shell.
There is a single large posterior adductor muscle at the center of the specimen which is
comprised of smooth and striated adductor muscles.

Discussion
Perna viridis and Geloina expansa both possess two siphons, the inhalant, and
exhalant siphons. Water is taken up by the large inhalant siphon and pumped through
the gill filaments by a current caused by the beating of cilia found on gills. The bands of
lateral cilia that run along the sides of the gill filaments will then entrap the food-laden
water and organic food particles in mucus, then move them along the gill from the
posterior to the anterior of the bivalves where the labial palps are. The layered gills
increase the surface area. The exhalant siphon acts as an exit for the remaining water
and sediment particles to be pumped out. The labial palps regulate the food entering the
mouth and all rejected food particles are discharged as psuedofaeces. (Rajagopal et
al., 2006). The labial palps in both bivalve species are large and elongated like other
suspension feeders (Morton & Machado, 2019).
Perna viridis shells narrow at the anterior end and the anterior adductor muscle is
absent, this makes the anterior end narrower and allows it to wedge into crevices.
(Morton, 2022). As a result, the siphons are at the posterior end facing where the waves
crash on the rocks Perna viridis attaches to. Its siphons are short as it does not burrow
in sediment (Bales & Venable, 2007). Because Perna viridis do not burrow, its foot is
smaller and tongue-shaped to allow for vertical movements to climb rocks (Lalla, 2015).
The byssus threads are used to anchor themselves onto the rocks or hard surfaces, so
they can remain sessile during high flow or prevent them from getting washed away by
waves. Saccostrea also rests on rocky surfaces on their flatter valves. Saccostrea freely
swim around as juveniles before settling on a spot permanently for the remainder of
their lives, that is why they lack siphons, foot, and byssus as they do not need to move
around (Britannica, 2022

Geloina expansa and Ensis are both sediment-burrowing bivalves. Geloina expansa
have a large and hatchet-like foot, this makes it easier to dig into the sediment. Their
siphons help them to filter feed while remaining buried, however, the specimen
dissected in this experiment has a shorter than average siphon. It is likely due to
Geloina expansa starting to prefer burrowing shallower than it normally would. This
could be from the fact that as they have thick shells and are less prone to predation by
weaker predators unable to break them, there is less need to burrow deeper (Morton,
2022). Another likely reason is the possibility of fishes nipping off siphons so Geloina
expansa would be forced to burrow less deep and be easier to prey on (Cledón &
Nuñez, 2010). Contrastingly Ensis have thin shells and so burrow deeper and have
longer siphons and feet. The Ensis shell is long and laterally with the anterior side being
narrower and the posterior side being wider, which makes it a quick burrower as it can
still be dug up (Morton, 2022). Pectin is a free-swimming bivalve that inhabits open
offshore waters. As such it does not require siphons or foot. Pectin have smooth and
striated adductor muscles that allow for fast adduction, with the help of the wings and
the streamlined shell shape, water is propelled past the shell hinge and Pectin is
propelled (Morton, 2022). The ridges make the shell stronger.

Conclusion
The bivalves have evolved to be very efficient at surviving and thriving in their habitat
and their habits. The reasoning for the shortened siphons of Geloina expansa is still
unknown. Further research needs to be conducted.
 Supplementary

Table 1 shows the comparison of the 5 species of bivalves’ habits, habitats,

and morphology.

Table 1 Habit, habitat and morphology of Perna viridis, Saccostrea, Geloina expansa,
Ensis and Pectin.
Perna Saccostrea Geloina Ensis Pectin
expansa
Rock mussel Rock oyster Brackish Razor clam Epibenthic
water clam scallop
Habit Anchors to Anchors to Burrows into Burrows into Swims freely
rocky rocky surfaces sediments sediments
surfaces
Habitat Shallow Intertidal Mangrove Intertidal or Offshore
coastal zones swamps subtidal waters
regions zones of bays
or estuaries
Morphology
Shell shape Elongated Long, Trigonal, Elongated, Fan-shaped,
triangular- irregular, flat, Rounded narrow, and wings
ovate and rounded convex
valve
Shell thickness Thick Thick Thick Thin Thin
Shell surface Ridges Ridges Ridges Ridges Ridges
Shell valves Symmetrical asymmetrical Symmetrical Symmetrical Symmetrical
Hinge location (shell) Anterior end anterior end Dorsal end Anterior end Dorsal end
Interlocking Interlocking Interlocking Interlocking Interlocking Resilium
mechanism (shell) hinge teeth hinge teeth hinge teeth hinge teeth
Adductor muscle One on the One in the Two, one on One at the one, at the
(number and posterior side middle the posterior posterior end center closer
location) side and one to the
on the posterior side
anterior side
Foot Small Absent Large Large Absent
Siphons Short, Absent Shorter than Long, Absent
inhalent average, inhalant
fringed inhalent fringed
fringed
Mantle Mantle fusion Mantle fusion Mantle fusion Mantle fusion Mantle fusion
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