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Iguana protein issues

IGUANA EVOLUTION
Iguanas have evolved to fill a certain niche - that is, herbivory in arboreal tropical environments. They are successful, because there are few competitors (especially reptilian ones). This pressure to efficiently digest plant matter has led to a longer, more sophisticated digestive system to enable fermentation and breakdown of otherwise quite hard to deal with matter. However, the pressure has been there because of the abundance of food, and perhaps the lower energy requirements necessary to locate and acquire food sources. Take a pinch of salt, add several hundred thousand years, and you have a digestive system nicely adapted for herbivory (I won't say perfectly adapted, because perfection is a very subjective quality!).

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RESEARCH ON WILD IGUANAS
None of the published scientific field research on green iguana diets in the wild suggests that they take animal protein, but it does show that they derive all their protein (and other) requirements from plant materials in the diet. Anecdotal reports which seem to have made it into the literature suggest that juveniles have an increased propensity to take insects. There is the odd report that iguanas also take bird eggs and carrion - there's even less evidence for either of these. The nutritional research which has been done (examining wild iguana gut contents) shows no evidence of animal protein, although if it were present it would be in such low amounts so as not to be detected by such a study. So, the natural diet of green iguanas does not include any animal protein. This doesn't discount the notion that they might accidentally ingest the odd insect together with a flower head, or whatever, but I hardly consider this to be anything significant... in the same way that cows don't eat animal protein, but they're not going to wrinkle their nose and lightly brush stray bugs from their juicy grass stems. My conclusion here is that they are strict herbivores under normal circumstances.

ADAPTATIONS TO HERBIVORY
Take a look at iguana morphology and physiology. Iguanas are not equipped to hunt down and digest animal prey. Their digestive system employs a hindgut fermentation process, which is to break down plant matter, cell walls and carbohydrate. This isn't to say they couldn't digest animal protein, but just that they're optimised for plant matter. They also spend their entire time in the wild living on their food plants. Why spend time and energy hunting animals when plant food is all around them? So, my conclusion that they are strict herbivores still stands.

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EXCEPTIONS TO THE RULE?
Feral green iguanas (e.g. Puerto Rico) have been reported to take insects more frequently that "incidental" would imply. Captive iguanas will also readily take insects, mice and other sources of animal protein if offered them (mice are often taken by iguanas that would happily remove your finger at the same time - almost giving the impression that they would eat anything presented to them). Some people seem to have almost converted their iguanas into carnivores. However, iguanas are nothing if not opportunistic. It seems that under conditions of stress and poor food availability (and that includes feral populations, which are mal-adapted to the local flora), they will resort to using additional food sources until the situation improves. This suggests that even in their native habitat they might resort to limited omnivory if that becomes necessary for survival. Iguanas in captivity are nothing if not stressed, and offered poor quality diets on the whole. Therefore, eating insects, mice or whatever doesn't seem unreasonable. Iguanas will happily eat their substrate if nothing else presents itself. Herbivorous animals have been known to eat carrion, including their own kind, when presented with no other food sources (e.g. cows and sheep have been reported to do this). Given this evidence, though, it seems that eating animal protein is a "last resort" - i.e. it's a poorer choice of diet. Also, in the wild, if a normally herbivorous iguana is forced to resort to eating some animal protein, the change will be temporary until better food options present themselves.

CONSEQUENCES OF HIGH PROTEIN INTAKE
Iguanas, and any animals for that matter, grow very quickly with increased amounts of protein. If you're interested in fast growth rates, usually when the animals are harvested for food, then increase their protein intake. As animal protein is better than plant protein for this, then feeding herbivorous animals animal protein will increase growth rate. This is why there is so much controversy over BSE in cows in the UK - beef cattle are pumped full of animal protein (including spinal tissue, in this case) so they grow quickly. However, the farmer is not interested in long-term health - the animals are culled at an early age, with the exception of breeding animals which recieve a different diet anyway. Dairy cattle are fed lower amounts of animal protein. Long term health is affected by excess protein, however. Veterinary evidence shows that excess animal protein in iguanas leads to gout, dehydration, metabolic bone disease, renal failure, kidney stones etc... and is one of the biggest causes of relatively young iguana mortality. The reason these poor diets (i.e. high percentages of animal protein) exist in the older literature is that they were formulated from farm-raising practises of green iguanas. So, iguanas show good growth rates, but the longterm effects severely shorten longevity.

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SPECIFIC EFFECTS OF ANIMAL PROTEIN ON IGUANAS
Animal protein contains components which iguana digestive physiology can't deal with in large amounts (e.g. fats, different purine/pyramidine ratios, different amino acid ratios). This also very likely unbalances the natural gut fauna the iguana possesses in the hindgut (as has been shown in studies on other herbivores), reducing digestion efficiency of plant matter which should make up the majority of the diet anyway. Iguanas, being herbivores, are physiologically adapted to process specific kinds of dietary protein - i.e. those found in plant materials. While animal protein can be metabolised, it literally overwhelms the system in excess (and it doesn't take much to cause an excess - although we don't know exactly how much). Nucleic acids in the foods are degraded to nucleotides and ultimately pyrimidine and purine bases - more are synthesised from amino acids in the liver. Any of these which are not used by the body are broken down into uric acid (or allantoin in some mammals, reptiles, fish). With animal proteins, green iguanas end up with an excess of uric acid in the body fluids, and over time this crystallises out - so uric acid crystals form in tissues throughout the body, impairing their function and leading to serious problems.

DO SAFE LEVELS EXIST?
There's no doubt that excess animal protein kills iguanas in the longterm. However, that's not the question. The question is, is there a certain amount of animal protein (very low percentage of daily intake) which will either not harm the animal, or perhaps even be beneficial because of its high protein content? Hindgut fermentation works best in larger animals (more efficient per unit volume of food), but what about hatchling iguanas which are much smaller? Would perhaps the odd insect lead to increased growth with no obvious health problems until the hindgut area grew large enough to be able to reach a greater operating efficiency? (i.e. they're not utilising all the purines/pyramidines in their diet, so animal protein doesn't cause uric acid concentrations in the blood to go up significantly because not all the purines/pyramidines are being utilised)

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BUT ISN'T ANIMAL PROTEIN REQUIRED FOR...?
It is quite commonly believed that animal protein is the only way an iguana can get any or enough calcium in it's diet (because animal bones contain calcium). This is incorrect - calcium is present in varying quantities in most plants, but still many iguanas suffer from calcium deficiency which leads to metabolic bone disease. In order to prevent these symptoms, a balanced diet needs to be created which contains sufficient calcium and sufficient vitamin D3 (which may be present in the diet, supplements, or more usefully through synthesis in the skin by the action of UVB wavelengths in sunlight). At the same time, the inhibiting effects on calcium metabolism of phytates, oxalates, acid and fat must be minimised. Powdered calcium powder helps to supplement a diet which might otherwise be low in this important element. It is also sometimes believed that animal protein is the only way an iguana can get any protein, period! This simply isn't true - plants also contain proteins, but with different ratios of amino acids to animal tissue (the former of which the iguana digestive system has evolved to cope with).

FURTHER SPECULATION ON NATURAL VARIATION
Ranging from Mexico to Brazil as they do, iguanas come from a diverse array of habitat types and it's very likely, given the morphological and behavioural varitations that exist, that similar variations in dietary preferences also exist. Perhaps some populations do select limited amounts of animal protein, and perhaps these animals do have lower life expectancies. We simply don't know this yet. There are clues: some iguanas appear to reach much smaller adult sizes than their counterparts. It's one of my speculations that these animals may be adapted to a poorer diet (caused by adaptation to the different foods available in another habitat), and even if presented with a richer diet don't grow as quickly as those from other populations. Although Iguana iguana is one species, there are many different races which can often become available in the pet trade. Remember, though, that captive farmed iguanas (which make up by far the majority of animals available) are coming from a limited number of breeding operations, which are harvesting eggs/raising animals from the same area, and therefore diversity in captive populations may be more limited. I'm a little wary, though, of suggesting a perfect diet for all iguanas (from Mexico to Brazil) - it doesn't make any sense that they would all benefit equally from exactly the same diet given that they don't live in the same environments, but until we learn more (and we haven't yet) the best we can do it provide diets which we know are beneficial to the widest number of animals.

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GENERAL CONCLUSIONS
Iguanas are herbivores. However, like adding leaded fuel to an unleaded car, you can still throw other things through the digestive system, and it will still try and do what digestive systems are good at - digest it! A small amount of leaded in an unleaded car might make it hiccup at little, but no lasting harm. In captivity, though, people put too much leaded fuel in and destroy the engine of their iguana over time. The whole digestive system can cope with much of the contents of animal protein, but because the rest of the body isn't used to dealing with this kind of food source (inc. high protein, fat etc...), things get thrown out of balance if there's too much of it (this doesn't normally happen under natural conditions, otherwise selection pressure would tend to lead to more omnivorous animals).

Those who think that green iguanas should for some reason be fed animal protein are not considering their natural physiology or behaviour. They know that animal protein (i.e. large amounts of protein) will result in additional growth. They're right - just look at beef cattle. However, much of this information has come from captive farming operations where animals were intentionally grown quickly for either reintroduction programs or for food purposes. In the former, the longterm negative effects were not known, although if it was enough to allow several breeding generations of wild animals then its purpose would have been satisfied. In the latter, who cares if an animal will die at 5 years if it's slaughtered for lots of meat at 3 years?

Interestingly, many of those formulating diets for captive iguanas for the pet industry still include animal protein, because there's a belief that there must be some level at which animal protein benefits growth without signficantly affecting longterm health. I think this view is erroneous, particularly as they still have little idea what kind of protein levels are harmful - taking a chance, if you like, but if the food makes your ig grow, people are going to buy it, right? Perhaps this is a factor - if someone buys product A that includes animal protein, their iguana will appear to grow faster. But, if they try product B because they hear it has no animal protein, but their iguana's growth slows down a little, they might go back to product A. Cynical, but probably not without merit! If someone does decent longterm research and proves my view wrong, then fine, but until they do my opinion stands. After all, a good artificial diet is theoretically possible.

My overall conclusion to this is that no, we don't know it all. Far from it. However, everything that we do know so far tells us to be very cautious. I'm sure the odd bit of chicken, insect or whatever on very rare occasions will do the iguana no harm. However, I really don't see why this should be necessary, and I can't understand the hangups about missing it out altogether. It's almost as though people feel that iguanas must have a little animal protein every so often, otherwise they might be missing out on something. I get the impression others tout animal protein just because everyone else says don't bother with it. When the idea of using animal protein is so entwined within the literature of even fairly modern books, it's hard to avoid this question - it keeps coming up time and time again. This is good, because every time it comes up, we hopefully know a little more about how the processes work. At the moment, however, we don't know enough, and so my advice to people is always not to include it - not because a tiny bit might not be beneficial, but because too much is definitely bad, and we don't know how much is too much! People who use animal protein are, in my opinion, playing dice with the health of their iguana (longterm), and I don't think it's worth it - given that we know a totally herbivorous diet is very healthy - but a good totally herbivorous diet, that is! A bad diet, period, will kill your iguana. But I think the best diet is totally herbivorous, until I see any convincing evidence to the contrary, or firm data to show how much animal protein can be safely utilised (if any).

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FURTHER READING
Donoghue, S. (1996). Nutrition of the Green Iguana (Iguana iguana). In: Proceedings of the Third Annual Conference of the Association of the Reptilian and Amphibian Veterinarians (Frahm, M.W. ed.). ARAV. pp. 99-106

Donoghue, S. & Langenberg, J. (1996). Nutrition. In: Reptile Medicine and Surgery (Mader, D.R. ed.). W.B. Saunders Company, Philadelphia. pp. 148-174

Marken Lichtenbelt, W.D. van. (1993). Optimal foraging of a herbivorous lizard, the green iguana, in a seasonal environment. Oecologia 95: 246-256

Rand, A.S., Dugan, B.A., Monteza, H. & Vianda, D. (1990). The diet of a generalized folivore: Iguana iguana in Panama. J. Herpetol. 24: 211-214

Van Devender, W. (1982). Growth and ecology of spiny-tailed and green iguanas in Costa Rica, with comments on the evolution of herbivory and large body size. In: Iguanas of the World: their Behaviour, Ecology & Conservation (Burghardt, G.M. & Rand, A.S., eds.), Noyes Publ., Park Ridge, N.J. pp. 162-183