Invertebrate anatomy

KU022 Physiology of Aquatic organism Bent Vismann

cnidarians

bivalves flatworms nemerteans gastropoda pygnogonida

ctenophores

chelicerates bryozoans pogonophorans sponges cephalopods annelids crustaceans Echinoderms ascidians

cnidarians

bivalves flatworms nemerteans gastropoda

ctenophores chelicerates bryozoans pogonophorans sponges cephalopods annelids crustaceans Echinoderms ascidians

TAXONOMY
Phyllum: Mollusca

Gastropoda

Bivalvia

Cephalopoda

Protobranchia

Lamellibranchia

Some commercially exploited species

Venus clams (Venus, Mercenaria, Cyclina sp.)

Pacific Oyster (Crassostrea gigas) European oyster (Ostrea edulis) Edible cockle (Cerastoderma sp.) Scallops (Pectinidae) (Pecten maximus)

Jackknife clam (Ensis sp.)

Blue mussel (Mytilus edulis)

Epifaunal bivalves

Infaunal bivalves

Siphon

Geoduck (Panope generosa)

App. 5 kg and siphon up to 2 meter

Just an amorphous slimy lump with no distinct feature?

aorta ventricle auricle visceral artery nephrostome

pericardium intestine

anus organs

kidney

longitudinal vein

gills Mantle artery

mantle

Adult blue mussels sperm

egg spat In e.g. oysters this takes place at the gills (i.e., it is the veliger larvae that comes out from the adult bivalve) Pediveliger larvae Trochophore larvae Veliger larvae

embryo

Heart

Digestive gland

stomach Efferent cavity

Afferent cavity mantle

Mantle cavity

W-shaped gill

Laterofrontal cilia Frontal cilia Branchial vessels Ciliary junctions

Lateral cilia

Interlamellar junctions

Food grove

How to measure filtration?

FR: Filtration rate (or Pumping rate) The water volume pumped per time unit

CR: Clearance rate The water volume cleared for particles per time unit

RE: Retention efficiency The gills efficiency to retain particles CR = FR * RE When RE = 1 CR = FR

In order to know how many algae is present (or removed) we need to quantify the algae

Microscope and counting chamber

Cells ml-1 Cells ml-1

Coulter counter (electric registration of particles sucked through an orifice) Spectrophotometer (extraction of chlorophyll) g Chla ml-1

Fluorometer (excitation of chlorophyll and measuring the fluorescence)

Volt

Now turning to the bivalve A bivalve is not just a bivalve; i.e., size matters for any given physiological activity Relation between size and shape
Soft part (g) 2.5 2 1.5 1 0.5 0 0 10 20 30 40 50 60 70 Shell length (mm)

Allometric equation: W L3

Y=

aXb L W1/3

W = 3.3 * 10-6 L3.16

Mhlenberg & Riisgrd 1979

12 10 Clearance (l h )
-1

8 6 4 2 0 0 10

Cl

L2

Cl = 0.0012 L2.14

20

30

40

50

60

70

80

Kirboe & Mhlenberg 1981

Shell length (mm)

Cl L2
Clearance (l h )

14 12

and when L W1/3 Cl (W1/3)2 W2/3 W0.67

10 8 6 4 2 0 0 0.5 1 1.5 2 2.5 Soft parts (g)

-1

Cl = 7.45 W0.66

Jones et al. 1992

So, when measuring clearance it is important to realize that the clearance rate is dependent on the bivalve being measured. In other words, the clearance rate must be given in a form, which can be related to other studies. Also remember the same holds true for experimental conditions

Clearance (l h g )

20 18 16 14 12 10 8 6 4 2 0 0 0.5 1 1.5 2 2.5 Soft parts (g)

-1

-1

Closed system
4 Algae (1000 cells ml )
-1

3.5 3 2.5 2 1.5 1 0.5 0 0 10 20

Algae = a *e(mt)

Mussels pumping at a constant rate are placed in a tank with presence of algae This one can verify by a linear regression analysis

30 Time (min)

40

50

60

4 Ln(algae) (1000 cells ml )

-1

3.5 3 2.5 2 1.5 1 0.5 0 0 10 20

Ln(Algae) = ln(a) + mt

30 Time (min)

40

50

60

4 Algae (1000 cells ml )

-1

Closed system Algae = a *e(mt) Because the decrease in algae is exponential the ratio ln(algae0/algaet) is constant and describing the degree of decrease (i.e., whatever time period used on the graph the result will be the same)

3.5 3 2.5 2 1.5 1 0.5 0 0 10 20

30 Time (min)

40

50

60

Clearance = V/(n*t) * ln(algae0/algaet) V = volume 5 l); n = number of mussels (or biomass (4 g)); algae0 and algaet = algae concentration at time 0 and t, respectively Clearance = 5/(4*30) * ln(2.2062/0.5468) = 0.058 l min-1 g-1 = 3.48 l h-1 g-1

Weight specific clearance

Flow through system C1 C2 dC0/dt = F/V * (C1-C2) 1/V * C0 *Cl In steady state: Cl = F * (C1-C2)/C0 However C0 is difficult to measure, so a stirrer is applied: C0 = C2 In steady state: Cl = F * (C1-C2)/C2 If you use this equation without stirrer erroneous results might be obtained

F = flow rate V = Volume Cl = clearance

C0

4000 3500 Algae (cells ml )

-1

C1

3000 2500 2000 1500 1000 500 0 0 10 20 30 Time (min) 40 50 60

C2

In steady state: Cl = F * (C1-C2)/C2 = 0.174 * (3500-1500)/1500 = 0.232 l min-1 F = 174 ml min-1 W=4g = 13.92 l h-1 = 3.48 l h-1 g-1

Weight specific clearance

basic experimental set-up

Data on disk

Flow-trough fluorometer input Remote controlled peristaltic Pump

Analog input Pump

Interface output

Flow-trough Pump fluorometer

The set-up can be used in two modes: A) Continously clearance rate B) Intermittent clearance rate

Lo g

fu nc tio

Algae stock solution

Closed loop Control Output (on/off)

PC & Labtech Notebook

4000 3500 Algae (cells ml )

-1

3000 2500 2000 1500 1000 500 0 0 10 20 30 Time (min) 40 50 60

Cl = 1/(n * t) * (vp * tp* C1)/ C2 Where n = number of mussels (or biomass); t = time; vp = dosing pump rate; tp = dosing pump active; C1 = algae concentration in stock solution; C2 = algae concentration in aquaria

4000 3500 Algae (cells ml )

-1

3000 2500 2000 1500 1000 500 0 0 10 20 30 Time (min) 40 50 60

Cl = V/(n*t) * ln(C0/Ct) V = volume; n = number of mussels (or biomass); C0 and Ct = algae concentration at time 0 and t, respectively

Particles cleared

I PF R F P U

Particles retained by the gills can either be ingested (I) or rejected as pseudofaeces (PF) A part of the ingested food is not absorbed by the bivalve but is excreted as faeces (F) A part of the absorbed food is excreted as urine, mucus mm. (U) The remaining part of the absorbed food is called the assimilated food and is used for growth = Production (P). In connection with growth and activity some of the energy contained in the assimilated food is lost through respiration (R) Energy balance: I = P + R + U + F

P = I (R + U + F)

Max clearance at 2000 cells ml-1

AE = (I - (U + F))/I

Sejr et al. 2004

Energy: Algae = 1.75 J cell-1; O2 = 14 J mg-1; NH4 = 0.025 J g-1

I = Cl * [algae] P = I (R + U + F)
3.5 3 Energy (Joule h ) 2.5 2 1.5 1 0.5 0 -0.5 0 5000
-1

AE = (I - (U + F))/I

P = (I * AE) - R

Optimal production 6000 cells ml-1

ingestion Respiration Excretion production

In situ P
10000 15000 20000 25000
-1

30000

Algae concentration (cells ml )

Zero production at app. <300 cells ml-1

Zero production at app. >21000 cells ml-1

Crustaceans

Phyllum: Arthropoda Phyllum: Arthropoda Trilobitomorpha Trilobitomorpha Mandibulata Mandibulata Chelicerate Chelicerate Arachnida Arachnida Pycnogonida Pycnogonida

Chilopoda Chilopoda

Crustacea Crustacea

Insecta Insecta

Diplopoda Diplopoda Euphausiacea Euphausiacea Eucarida Eucarida Decapoda Decapoda

Copepoda Copepoda

Cirripedia Malacostraca Cirripedia Malacostraca

Mysidacea Mysidacea

Cummacea Cummacea

Isopoda Isopoda

Amphipoda Amphipoda

Telson

Cervical Abdomen Carapace groove Maxilliped Eye

Rostrum 1. antenna 2. antenna

Uropods

Pleopods

4. walking leg

1. walking leg

Cheliped

Intestine Abdominal extensor Dorsal abdominal artery

Ovary

Opthalmic artery Cerebral ganglia

Pericardial Antennary Ventral artery Nerve cord sinus Heart

Abdominal flexor

Ganglion Ventral thoracic artery

Digestive Cardiac gland stomach

Green gland

Ventral abdominal artery

Esophagus Pyloric stomach Circumesophagus Sternal connective artery

Hemocoel

artery

Heart Gills