Birth and hatching: Key events in the onset of awareness in the lamb and chick

Abstract

The neuroanatomical and neurophysiological development of the embryo and fetus and unique features of the physiological environment of the fetal brain, features which are lost at birth, support recent conclusions that under normal circumstances awareness (or consciousness) is probably not exhibited by the ovine embryo-fetus before birth and that it appears for the first time only after birth. However, there has apparently been no evaluation of whether or not similar mechanisms modulate awareness-related functions in domestic chicks before and after hatching. This comparative review, in seeking to rectify this, arrived at the following conclusions.

First, the neural apparatus of both lambs and chicks appears to be too immature to support any states resembling awareness during at least the first half of pregnancy or incubation. Second, electroencephalographic (EEG) activity, which evolves subsequently, shows that states of sleep-like unconsciousness are likely to be continuously present in lambs until after birth, and that such states at least predominate in chicks until after hatching. Third, as in fetal lambs, epochs of so-called ‘wakefulness’ previously reported in chick embryos do not seem likely to represent short periods of awareness in ovo. Fourth, several neurosuppressive mechanisms, with some unique features, also operate or have the potential to operate in chicks before hatching, but a dearth of published information currently hinders a full comparison with those demonstrated to operate in fetal lambs. Fifth, contradicting the intuitive perception that vocalisation pre-hatching by the chick indicates the presence of awareness, published evidence suggests that vocalisation before and during hatching occurs mostly during EEG states indicating sleep-like unconsciousness. Sixth, as seems to be the case for newborn lambs after birth, it is possible that demonstrable awareness may appear for the first time only after hatching in chicks, presumably through waning neurosuppression and burgeoning neuroactivation, but such awareness seems to take longer to manifest itself. However, additional research in chicks is recommended to further assess this suggestion. Particular attention should be given to the status of vocal interactions between hen and chick which begin several days before hatching, and to the operation of neurosuppressive and neuroactivating mechanisms throughout the last 40% of incubation and during and after hatching.

KEY WORDS:

• Behaviour
• electroencephalogram
• developmental neurobiology
• embryo
• fetus
• neuroactivators
• neurosuppressors
• onset of awareness
• unconsciousness

Acknowledgements

We thank Professors Don Broom (Cambridge University) and David Morton (University of Birmingham) for asking questions about chicks that stimulated us to prepare this review. We also very much appreciate the most helpful input received from Associate Professors Laura Bennet and Alistair Gunn (Auckland University), and from Associate Professor Roger Lentle, Dr Craig Johnson and, especially, Mr Bradley Siebert (Massey University). Likewise, constructively critical comments from five anonymous referees are very much appreciated. We gratefully acknowledge the Ministry of Agriculture and Forestry (New Zealand) for financially supporting our related investigations in lambs, and the Agricultural and Marketing Research and Development Trust for doctoral scholarship support for TJ Diesch.