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NEUROTRANSMITTERS IN NEMATODES 365<br />

approximately 300 neurons in A. suum. Sensory<br />

and motor neurons are concentrated at<br />

the head and the tail of the nematode. The<br />

rest, about 90 motor neurons, are arranged in<br />

repeated segments along the length of the body;<br />

they supply the innervation to the somatic<br />

muscle cells. Figure 15.8 illustrates a segment<br />

of the dorsal and ventral nerve cord with<br />

connecting commissures (three to the left and<br />

one to the right). Each segment has 11 motor<br />

neurons that are excitatory (cholinergic) or<br />

inhibitory (GABAergic). This layout is believed<br />

to be very similar in all species of nematode.<br />

Due to an elementary system of reciprocal<br />

innervation of the neural circuits, excitatory<br />

impulses to the ventral muscle are associated<br />

with inhibitory impulses to the dorsal muscle,<br />

and vice versa. This combination of signals<br />

results in contraction of one block of muscle<br />

whilst the opposing muscle mass relaxes, allowing<br />

the nematode to progress forwards or backwards<br />

with a wave-like motion.<br />

Ascaris physiology<br />

Membrane potential<br />

The membrane potential of Ascaris suum<br />

muscle is near 30 mV, and is relatively insensitive<br />

to changes in the ionic composition of<br />

FIGURE 15.8 A ‘segment’ of the dorsal and ventral nerve cord of Ascaris, with 11 motor neurons and six longitudinal<br />

neurons. There are three right-hand commissures and one left-hand commissure in each segment. The diagram<br />

represents one of seven repetitive segments found in Ascaris. The filled, •, cells are excitatory motoneurons<br />

with acetylcholine as the fast <strong>trans</strong>mitter and empty, O, cells the inhibitory motoneurons with GABA as the fast<br />

<strong>trans</strong>mitter. FaRP peptides may act as co-<strong>trans</strong>mitters. Each motoneuron is classified by the location of the cell<br />

body, and the shape and distribution of its processes. All cell bodies are found in the ventral cord. The excitatory,<br />

cholinergic outputs, , are shown, as are the inhibitory GABAergic ouputs, . The inputs, are also shown. Dorsal<br />

excitatory motoneurons: DE1, DE2, DE3. Ventral excitatory neurons: V1, V2. Dorsal inhibitory neurons: DI. Ventral<br />

inhibitory neurons: IV. V1 and V2 do not have commissures. All commissures are right-handed except DE3.<br />

BIOCHEMISTRY AND CELL BIOLOGY: HELMINTHS

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