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38 Behaviour in models<br />

In situ studies<br />

are required to test<br />

whether larvae orient<br />

Information about<br />

cues and detection<br />

thresholds is scarce<br />

The first step is to detect whether fishes orient or not. Such information<br />

can be provided by field studies involving the release of larvae<br />

followed by divers 90,109–111 , or in situ orientation chambers 112 .<br />

The second step involves testing experimentaly the ability of larvae<br />

to detect environemental cues 100,102,103,113–116 . The potential for detecting<br />

a cue can be demonstrated in the laboratory by testing the preference<br />

of larvae toward a given environmental signal for example (e.g. coastal<br />

vs. oceanic water, reef sounds vs. random sound). Which cue is actually<br />

used in situ is currently unknown, due to the lack of experimental<br />

means of testing cues separately in the pelagic environment.<br />

The last step would be to describe thresholds for detection. This<br />

directly relates to the spatial scale over which cues can be detected<br />

and used for orientation. Knowledge in this regard is currently mostly<br />

lacking and is difficult to obtain, yet this is essential information for<br />

incorporation into models.<br />

1.5.4 Suggested implementation<br />

Orientation as a<br />

parameter or an<br />

emergent property<br />

The implementation of orientation is closely associated with that of<br />

swimming (both horizontal and vertical): orientation is simply a choice<br />

among the set of possible swimming vectors. Once again, two approaches<br />

can be taken: (1) behavioural rules in response to the environment<br />

can be defined a priori, based on observations and experimental<br />

work; (2) these behavioural rules can emerge from the model by defining<br />

the set of possible swimming vectors, a biologically sensible “goal” for<br />

larvae (e.g. settlement), and letting an algorithm choose the suite of<br />

decisions to achieve this goal (see Irisson et al. 117 for an example of the<br />

use of stochastic dynamic programming to solve such an optimisation<br />

problem).<br />

In both cases, orientation is a function which associates a swimming<br />

decision to the current state of a particle, such as<br />

f : state × time × environment → swimming(speed,direction)<br />

The amount of detail in the orientation behaviour is determined by<br />

what is incorporated in each of the left hand side variables. In the most<br />

simple model, when orientation is observed but the clues are unknown,<br />

orientation depends only on the position of larvae and on time. When<br />

responses to sensory clues are involved, the environment can include<br />

temperature, food, predators, current fields, land associated chemical<br />

concentrations, sun orientation, etc. If some kind of energy budget is<br />

present, the state of larvae can also encompass energetic reserves. This<br />

formalisation is therefore very scalable.

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