I should like to highlight the excellent article on "Neuroethology" presented by Günther K.H. Zupanc. It perfectly fits the high quality standards of Scholarpedia both in style and contents. Zupanc's contribution is perfect as it stands, so that it becomes difficult for the reviewer A to suggest improvements. I have proposed three additions to the text, (i) referring to the interactive structure of behavior motivations investigated by E.v.Holst and coworker, (ii) suggesting a summary of key principles, and (iii) providing a short note on the interactive nature between pretectal and tectal feature-analyzing systems. Again, I emphasize that these are just a few suggestions which are on the decision of Dr. Zupanc to consider. I have incorporated those suggestions in the article in Scholarpedia under the "edit" links.
RESPONSE BY GÜNTHER K.H. ZUPANC TO SUGGESTIONS MADE BY REVIEWER A: I am grateful for the suggestions and minor edits made by Reviewer A, and I am delighted to accept these changes.
RESPONSE BY REVIEWER A: I herewith confirm that the modifications by GÜNTHER K.H. ZUPANC in response to the suggestions made by the reviewer address reviewer's concern to accepting this article. From reviewer's point of view (09-15-2010) there is no significant reason for any substantial change in the article.
RESPONSE BY REVIEWER A to AUTHOR’S revision upon advice of REVIEWER B (cf. “minor comment” No.6). I strongly recommend not rejecting the photograph of Theodore Holmes Bullock from the Author’s first version. This is a matter of style. It is commonly accepted in the science community that Theodore Holmes Bullock is the main founding father of Neuroethology.
In his draft Scholarpedia entry on “Neuroethology”, Zupanc provides an overview of this field with an emphasis on its history, the choice of model systems, and a brief description of three classic neuroethological model systems. Available textbooks, relevant journals, and the International Society for Neuroethology (ISN) are briefly reviewed as well.
There is no doubt that the author is highly qualified to write this entry into Scholarpedia. And while the current draft is well written and touches on several of the fundamental aspects of neuroethology, it does fall short in several ways. In the following, I propose ways in which these short-comings can be addressed, so that this entry can serve as a starting point for the interested student or researcher and hopefully get the reader excited about our field.
1) In my mind, the first major section should tell the (uninitiated) reader what neuroethology is (the study of the neural basis of natural behavior), what its goals are (including the key principles discussed), and how it is situated in modern biology (e.g., how it relates to neighboring fields such as behavioral neuroendicrinology, behavioral ecology, behavioral genetics, etc.).
RESPONSE BY GÜNTHER K.H. ZUPANC: As suggested by the reviewer, the first section, which defines the term 'neuroethology', has been expanded.
2) As it stands now, a reader could get the impression that neuroethology is a discipline of the past. While key principles are discussed, this is done from a historical viewpoint (which is fine, just not enough), and only one of the references cited is from the 21st century (as if nothing relevant has happened in the past 10 years).
RESPONSE BY GÜNTHER K.H. ZUPANC: In the revised version of the text, six references cited are from the 21st century. Together with major changes in the text, including addition of an 'Outlook' section, the entry not only presents an overview of the past, but should also leave the reader with the impression that neuroethology offers exciting opportunities for future research.
3) It appears that all neuroethologists are neurophysiologists, which is obviously not the case. A more balanced treatment would include psychophysical, molecular, and evolutionary approaches as well as the historical contributions arising from more theoretical and analytical approaches (e.g., biological cybernetics).
RESPONSE BY GÜNTHER K.H. ZUPANC: Text has been added to show how concepts from disciplines besides neurophysiology have influenced the development of neuroethology. These disciplines include neuroanatomy (see ‘History and key principles’, 5th paragraph), biological cybernetic (see ‘History and key principles’, 6th paragraph), molecular biology (‘see Outlook’), and evolutionary oriented areas of research (see ‘Classical model systems in neuroethology’, 1st paragraph).
4) A short section on where the field stands today and an outlook to future developments would be very helpful. I find it striking that the author contrasts neuroethology with “molecularly oriented biological disciplines”. Does that mean he does not consider modern molecular approaches towards understanding the neural basis of natural behavior to be part of the field?
RESPONSE BY GÜNTHER K.H. ZUPANC: A section entitled 'Outlook' has been added. In this section, the impact of molecular biology and genetics on the future development of neuroethology is clearly emphasized.
5) Also, an increasing number of neuroethologists apply modern evolutionary concepts, thus being truly “comparative”. The author never explains what he means with “comparative approaches”. Only because someone’s work utilizes a non-biomedical model system does not make it comparative.
RESPONSE BY GÜNTHER K.H. ZUPANC: The text addressing the importance of comparative approaches and evolutionary concepts has been rewritten and expanded. Examples of investigations on both invertebrates and vertebrates have been added (see 'Classical model systems in neuroethology', 1st paragraph).
6) The importance of invertebrate model systems in neuroethology should be emphasized more. Except for the stomatogastric ganglion, this vast area is completely ignored.
RESPONSE BY GÜNTHER K.H. ZUPANC: In the revised version of the entry, examples of investigations on invertebrate model systems have been added to the text in a number of instances to provide a more balanced view of the relative importance of vertebrates versus invertebrates for neuroethological research. For example, a discussion of the brain stimulation work of Huber on crickets has been added; the work of Hassenstein and Reichardt on the beetle Chlorophanus and the housefly Musca is described; the comparative study of Elliott and Susswein (2002) on properties of the neural networks of gastropod molluscs is mentioned; the work on the neural basis of acoustic communication in crickets is presented as a classical example of neuroethological research; and the identification by Konopka and Benzer (1971) of a clock gene in Drosophila is mentioned.
7) I suggest expanding the list of classical model systems somewhat. Important examples include acoustic communication in frogs and crickets; echolocation in bats; escape behavior in cockroach and teleost fish; etc.
RESPONSE BY GÜNTHER K.H. ZUPANC: The list of classical model systems has been expanded by adding a description of the classical work of Franz Huber and associates on the neural basis of acoustic communication in crickets (see 'Classical model systems in neuroethology', 2nd bullet point).
Minor comments: 1) The author mentions Erich von Holst’s important brain stimulation work, but neglects to discuss earlier work by Franz Huber on crickets, which really was the first application of this methodology in a neuroethological context. In fact, Huber went to Hess's lab to learn this technique. This oversight exemplifies the lack of a more balanced discussion of vertebrate and invertebrate work.
RESPONSE BY GÜNTHER K.H. ZUPANC: As suggested by the reviewer, a description of the brain stimulation work of Franz Huber has been added (see 'History and key principles', 3rd paragraph).
2) The concept for central pattern generators predates the field of neuroethology by several decades. The way work on CPG is presented here is thus somewhat misleading.
RESPONSE BY GÜNTHER K.H. ZUPANC: The discussion of the importance of neuroethological studies for the understanding of central pattern generators has been modified (see 'History and key principles', 7th paragraph).
3) Animals do not produce “fictive behaviors”; rather, in reduced preparations these are phenomena that approximate the real behavior and can be measured reasonably well.
RESPONSE BY GÜNTHER K.H. ZUPANC: The discussion of the importance of the analysis of fictive behavior for neuroethological research has been rephrased (see 'Choice of suitable model systems', 2nd paragraph).
4) The classic book by Camhi should be mentioned. It probably is the best book on neuroethology ever written, even though it is now out of date (as is Ewert’s)
RESPONSE BY GÜNTHER K.H. ZUPANC: As suggested by the reviewer, the book by Camhi is now mentioned. A scan of the cover page of his book has been added (see 'Textbooks', 1st paragraph and Figure 1).
5) Under the Professional Society heading, add brief info on the most recent and next meeting, as well as a reference/link to the (upcoming) Gordon Research Conference in Neuroethology.
RESPONSE BY GÜNTHER K.H. ZUPANC: As suggested by the reviewer, the history of past meetings, and information on the next meeting, of the International Society for Neuroethology have been added. Furthermore, reference/link to the Neuroethology: Behavior, Evolution & Neurobiology Gordon Research Conference has been added (see 'Professional Society and International Meetings').
6) If photographs of pioneers are included (which I consider unnecessary; links to their own entries are best), I don’t think it is appropriate to only show Ewert and Bullock.
RESPONSE BY GÜNTHER K.H. ZUPANC: The photographs of Ewert and Bullock have been removed. Neither of these two scholars has an entry in Scholarpedia.
I think the author has done a very good job of revising the entry on Neuroethology, and I accept the revision. Regarding the comment by Reviewer A to not remove the photo of Ted Bullock, I would encourage this reviewer to write an entry on this and maybe other eminent figures of our field. As indicated by the author, no such entries exist at this point.
User 4 (Editor ): What is Ethology?
In the article, it states: "By contrast, ethologists employ a whole-animal approach. Preferably, they study the behavior of an animal in its natural habitat, avoiding disturbance by the experimenter as much as possible. If investigations in the laboratory are performed, the animal is kept under conditions mimicking those in the natural habitat as closely as possible. Many of the behaviors studied are rather complex, often occurring in the context of social interactions between conspecifics."
I don't think that this is true. There is generally substantial manipulation and interference by the experimenter. Think of Tinbergen using colored sticks to mimic gull beaks or changing the length of a sword tail or finding the sign stimulus for a stickleback. Without manipulation, it's not really an experiment, it's just observation. --Pkatz 10:37, 14 October 2010 (CDT)
User 4 (Editor ): Suitable species
I don't agree with the statement, "The animal displaying such behaviors should be inexpensive and suitable for maintenance and breeding in the laboratory. Furthermore, the neural network underlying the behavior should be relatively simple in the sense that the nervous system consists of a rather small number of neurons, representing a minimal number of different classes of nerve cells."
I work on animals that are wild caught (Tritonia) and are not suitable for breeding in the lab. Michael Platt and Katarin Gothard consider themselves primate neuroethologists. Primates are not inexpensive and they don't have a small number of neurons. No mammal will fulfill the criterion of having a "simple" nervous system, but there is a substantial body of mammalian work that should not be excluded from neuroethology. --Pkatz 10:50, 14 October 2010 (CDT)