IV. TRACE INORGANICS

IV. TRACE INORGANICS

The field moves very slowly forward.Proving essentiality has proven more difficult with each new inclusion. The first two papers are not limnological, not even ecological. (Then again - maybe they are.) However, you draw the barriers, each is written by a legitimate leader in the field of trace element nutrition. It "gets me where I live" to acknowledge that most ecologists (that encompasses my fellow travelers in limnology) do not appreciate the significance of trace elements. They tend to mumble that most of them are around in the soil if they are terrestrial ecologists, and that most of them are around in the water if they are limnologists. It was a great comfort to note that in the chapter from his 1991 book, you read a selection from it (under eutrophication), Edmondson acknowledged that Se and Zn might be in short supply in some waters. But, even there, I have a feeling from conversations that that was, at the time, a tentative acceptance. He seemed comfortable concerning their actual influence in fw community structure only after I found a reasonable reference in a National Academy Report (he later sponsored a paper about our Zn/Se work to the academy - but he wanted to be convinced!).

The only influential ecologist whom I know who really felt strongly about the significance of trace elements was Hutchinson. Dr. Patrick, who has written about the trace elements on several occasions, does not think the time has yet come for emphasis. But she has published a new book (1996) and maybe (just maybe) she will try once again to get the limnological community to admit they cannot be ignored. She mentioned that intent just prior to publication. Unfortunately, Hutchinson (and Patrick, I suspect) felt that only time would make colleagues recognize their enormous omission. Hutchinson gave some of the trace elements their "due" in the Treatise on Limnology Vol. 1 (that was written in 1957 !!). But, there was then, and is now, an amazing lack of solid information concerning the trace element needs of any organisms, especially of any aquatic organisms. (You really should read the Treatise - but --- just come visit it some day.) The basic problem is that it is extraordinarily difficult to control trace elements in any test system.

Provasoli, usually identified as a "biological oceanographer interested in plankton nutrition", was absolutely sold on the notion that the trace elements are a critical force in plankton community structure. He lamented at length that the state of the art in chemical preparation made it almost impossible to properly test this hypothesis. At least everyone in the field of phycology is convinced that the trace elements limit growth in the open ocean. [Nitrogen limits in coastal {COASTAL} waters; phosphate is in relativel abundance]. But, Provasoli was continually frustrated in his efforts to prove that by the state-of-the-art of chemical purification. That same state-of-the-art today is, finally, about ready to support such work. (That is what we do in my lab.) Provasoli (who clearly thought as an ecologist) always referred to ecologists as "other", whereas Hutchinson considered them "self". BTW - the identification and quantification of trace elements (or anything) is not the same thing as determining what is actually available to a creature.

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Schrauzer, G. 1984. Introduction to Chapter 2. Biochemistry of the Essential Ultratrace Elements. (Series: Biochemistry of the Elements, Vol. 3):17-31. Plenum Press, N.Y. A leader in the field, offering a state-of-the-art overview. He, Mertz, and Forest Nielson are (I am fairly sure) the "grand old men" of the trace element business - all are active today (1999).

Mertz, W. 1981. The essential trace elements. Science 213:1332-1338. Just what constitutes proof of essentiality? For some organism other than a human? For a human? Some newly published "proof" is often hailed as THE proof of essentiality, even though some earlier "proof" was also hailed as THE proof. What is Mertz's concept of the "total dose-response range"? It should remind you of Parasalsus ("As the dose, so is the poison"). The Eskew, et al., paper from 1983, infra, is THE latest (that’s how slow progress has become, as of 2003) addition to the proofs of essentiality - it only covers plants (microbes, plants, animals are the needed categories) and it does not really offer a "kill". In 1998, the Nat. Acad. Food and Nutrition Board decided to count nickel as an essential for humans - even though hard "kill" proof has never emerged for plants or animals (only for Prokaryotes). Still, it comes closer than anything else in the last several DECADES (>40 yrs)to having been proved essential in the classical sense. The problem is dirt (contamination). There are a fair number of elements still suspected of being essential. A couple more than you might find in the Schrauzer paper - even though Schrauzer, himself, might well agree that they exist. He would avoid listing all the "possibles" because folks often misunderstand.

Eskew, D.; Welch, R., and Cary, E. 1983. Nickel: An essential micronutrient for legumes and possibly all higher plants. Science 222:621-623. A good example of the detailed proof offered when a scientist claims proof of essentiality. Think about the date. Do you realize how little we know "for sure"? Vide supra for the reason this paper is here.

Dr. Bartha (an emeritus member of this grad. program) is credited with identifying nickel as an essential nutrient. He did it with a bacterium. By the way Bartha killed the bacterium in conjunction with his proof. The best they could do for the plant-proof was to damage the leaves. No animal proof has yet been published. Strong suggestions abound concerning the need for nickel in animal metabolism, but no one has yet reported cleaning the contaminating nickel sufficiently out of a system to kill an animal. The reported damages are somewhat theoretical. How would you clean out the nickel in mouse feed? You will be regaled (oppressed) in class with a discussion of our nickel work to date. Especially, as it regards Fe. We managed to kill daphnids by clearing their world of nickel. That work is not yet published. The reason is important to your appreciation of the literature you encounter. Rest assured, you will hear it (and, probably many laments about controlling the test system.)

Patrick, R. 1978. Effects of trace metals in the aquatic ecosystem. American Scientist 66:185-191.

An unchallenged authority. This should be read carefully--not for the number of ppm which causes some change in dominance (these are just offered as concrete examples), but rather for the generalizations; e.g., the complement of species in a community is highly sensitive to changes in trace metals (true, not just for the first trophic level, but for all trophic levels). She is the only currently active and influential limnologist who is convinced that the trace metals matter to community structure - and she has, on several occasions, admonished me to forget about the T.E.'s, and get back to the study of trace organics only a couple of years ago. (She does not think the time is yet right for the TE.s when compared to trace organics.)

Eichhorn, G. 1975. Active sites of biological macromolecules and their interaction with heavy metals. Ecological Toxicity Research 6:123-142. You need not read the entire paper. I will ask you for a couple of examples of just how a metal ion can mess up a perfectly good molecule.(Just how does some metal ion get in there and do so much damage--there are a variety of ways discussed-take note of these) While you may not recall the details, you should not miss the obvious; i.e., those characteristics of the molecules in a living system which make them vulnerable to the intrusion of heavy metals. While you are at it, think of this material and consider why many metals have been shown to be both essential and toxic.

Provasoli, L., and I. Pintner. 1960. Artificial media for fresh-water algae: Problems and suggestions. The Ecology of Algae (Pymatuning Symposia). A lucid discussion of the problems one must consider when trying to satisfy the full complement of an organism's requirements. The easiest part is to include all the essential elements (just dig up a periodic table). But--how much is enough and how much is too much? This twenty + year old paper is still one of the best modern discussions of the topic. Most people in the field have not yet discovered most of the problems to which the authors allude. Remember presence does not equal availability.It is apparently easier to reinvent the wheel than it is to read any publication over 5 years in print. Read this one carefully. There is a second paper in next week's list which complements this one.

The following two papers are here because I felt like including them. I was serious, this really is what we do in the lab. Selenium, zinc and copper play an unusual role in toxicological work. That is not why we studied them. Actually, we studied them because a) our animals fell apart one summer, and b) because the concept of a "cheap shot" backfired.

Keating, K.I. and B.C. Dagbusan. 1984. The effect of selenium deficiency on cuticle integrity in the Cladocera (Crustacea).P.N.A.S.,U.S.A. 81:3433-3437.

Keating, K.I. and P.B. Caffrey. 1989. Selenium deficiency induced by zinc deprivation in a Crustacean. P.N.A.S., U.S.A. 86:6436-6440.

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Media lists can provide considerably more insight than might be initially obvious. See if you can tell the obvious difference between media for (what is, is not, included) -

- algae: autotrophs.

- animals: heterotrophs.

Type 1: Conklin and Provasoli, Moina macrocopa medium--for use when NO food organisms are fed.

Type 2: MS (in the selenium paper)--for use when food organisms are included in cultures.

Type 1 offers no control of trace inorganics.

Type 2 offers no control of trace organics.

The remaining papers are for rummaging. I believe that silica limitation plays an important role in plankton community structure and that it will ultimately play an important role in the development of controls for the most distasteful results of cultural eutrophication. Read what you will. If you really want some interesting material on this sort of thing, there is an overload of such in my lab/office/library/livingroom/familyroom/bedroom/kitchen/bath/cellar/attic/garage/spareroom/car/------