Defensive inking in Aplysia spp: multiple episodes of ink secretion and the adaptive use of a limited chemical resource.

The seahare Aplysia spp. extracts many of its defensive chemicals from its red seaweed diet, including its purple ink, which is an effective deterrent against predators such as anemones and crabs. It is believed that the inking behavior is a high-threshold, all-or-none fixed act that nearly completely depletes the seahare of its ink supply. If a seahare depletes its gland of ink, it must seek out a source of red seaweed and then feed for at least 2 days to replenish its ink supply. This suggests that the animal would not be able to deploy ink more than once in rapid succession in response to successive attacks from one or more predators. However, we found that Aplysia spp. can secrete ink in response to three or more successive stimulations with (i) anemone tentacles, (ii) a mechanical stimulus, consisting of grabbing and lifting the animal from the substratum, or (iii) a noxious electric shock. A spectro-photometric measure of ink secretion showed that only approximately 48 % of the gland's releasable ink reserves are deployed initially. Thus, deployment of this defensive chemical is not strictly all-or-nothing, although the trigger mechanism is. Moreover, the animal tends to secrete a relatively fixed proportion (30--50 %) of its available ink reserves even after its gland has been depleted to approximately half its initial content. Since an animal need only use a proportion of its ink reserves to deter an attacker effectively, the inking behavior is adaptive in its economical use of a limited resource.