Sunday, October 18, 2015

Thermodynamic equilibria of animals with envirrnment, Porter and Gates (1969)

In which two scientists endeavor to prove that animals do not, in fact, violate the laws of thermodynamics.

It is interesting to think of this paper as the authors sought to include it, demonstrating three concepts that were being explored in ecology at the time. Using mathematical models to better understand factors that might affect organisms in real life. Using physical models to try and recreate at least a simulacrum of the study organism and be able to analyze factors that affect them. Lastly, the idea of a "climate space" which we have been talking about through a lot of these paper. The idea behind this is the climate part that N-dimensional hypervolume that we consider when we are thinking about a niche.

The paper starts out talking about the idea of the climate space right away and gives us the first four dimensions of the hypervolume they will be considering: radiation, wind, air temperature, and humidity. These four factors are a pretty good approximation of climate. I am sure there are a number of other things that we could all come up with that might go in to climate but almost everything that goes into what a climate is comes from these four factors. These abiotic factors affect three factors they consider on the animal: body temperature, rate of moisture loss, and metabolic rate. They are analyzing the animals to be part of a steady state system and fully admit that this is not necessarily the case in the wild. But, the idea behind it is sound. Over a long period, the animal must be in a steady state. To not be in a steady state would mean that the animal was either cooling or warming over the long term. This is fairly contrary to what we know about animals, barring some age-based body temperature differentials.

These ideas about what goes into and out of an animal form this idea in the paper called the "energy budget" and this is just the idea that Energy IN = Energy OUT. But what they are really trying to do is to determine what the parts of these in and out equations. A quick rundown of the variable might be helpful.

Energy In

  • Metabolic Rate
  • Radiation Absorbed
Energy Out

  • Epsilon-sigma-Tr^4 is the energy lost by radiation of the animal
  • hc(Tr-Ta) is the energy lost by convection coefficient
  • Eex is the energy lost from expiration
  • Esw is the energy lost to sweating
  • C is the energy lost by conduction to substrates
  • W is the work done by the animal
The reset of this section is talking about how they obtained these values and decided on specific ones. It discusses some other factors (like color) that will have to go into calculating these values and discusses the effect of size on things like the convection coefficient.

In the next section, Heat Transfer Within The Animal, Porter and Gates go on to talk about a really important part of how animals interact with their environment, their adaptations to the environment around them. The are specifically concerned with the fat layer and the feather/fur layer that many animals have to help them adapt to different climates. Since they are treating all animals as cylinders, they look at the idea of concentric cylinders with the heat generating stuff in the middle most cylinder, a cylinder of fat around that, and a cylinder of feathers/fur on the outermost layer. These approximations give them a means to talk about how animals adapt to their environments to prevent or enhance heat loss.

This section leads into the next two sections which are more explanation of the equations. Specifically they talk about the energy that can be exchanged on the surface of the animal and the radiation that will be absorbed on the surface of the animal. These are both a function of the shape(cylinder)/size of the animal, and the energy exchange also has to do with the idea of the concentric cylinders that they are using as their animal model.

Finally they get in to their modeling of different animal's climate spaces. Using molds of animals as well as mathematical approaches, they took measurements of the energy absorbed by animals at different temperatures and amount of radiation absorbed. They admit that some things have to be known before their analysis can work and that animals can change some of these values about themselves, but they are still confident, and rightfully so, that their calculations and experimentation can show some valuable results. In their analysis they are able to find temperature ranges and sun radiation absorptions that would let an animal live in the temperatures that are within it physiological range. They do this with a number of animals (iguana, shrew, finch, pig, etc.) as well as some theoretical animals, showing that it works and can be extrapolated for other organisms.

In conclusion, they are defining their climate spaces as a three dimensional space of radiation absorbed, wind speed, and ambient temperature. While we know that there are definitely more factors, this is a really great step towards understanding the fundamental niche of an organism. This paper shows methodological advancements in a way that we haven't really seen before, using both mathematical models as well as physical models to test concepts. These two modeling methods are still used extensively today.

9 comments:

  1. As stated in Noah’s summary I thought these guys did a really nice job of expanding on the abiotic components of a niche. They took the concept of a fundamental niche to another level and tested it and defined it by specific climatic parameters. I think they were pretty thorough in this, but I thought the effect of water was somewhat one sided (I also may have missed something). They said a lot about dissipating heat with water but not much about the effect of being wet in cold temperatures and how that might affect survival temperature. The specific descriptions of animals were pretty wordy it seems like a figure might have been a better expression of this. They could have detailed unique characteristics or behaviors in the text and left temperature ranges to the figure.

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  2. I think this paper does a great job of detailing the factors that seem to have the most effect on a niche. While I may have not put them in the same terms that the authors did, I think they got the right ideas. I also find it so interesting that this paper aims to quantify the different anatomy parts of an animal (in general terms) in order to calculate how they each are affected by the abiotic factors. The variables that the authors came up with to quantify "energy out" seem to cover a lot, but I agree with Eric that the effect of water wasn't inclusive. Water plays a large part in animal survival in ways other than what they listed, including losing heat in extremely wet and cold temperatures. Overall, I liked this paper for what it did to show the factors included in the fundamental niche of an organism. I think incorporating modeling with a case study gives the experiment more credit and be more influential. It also appeals to a wider audience.

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  3. I thought this paper was somewhat difficult to get through. I again had to go back to Jim Brown's introduction to this section to get a better appreciation of the paper and what it added to the field of ecology. As already stated, this paper aimed to really flesh out the components of an individuals fundamental niche and how the specific anatomy of an organism interacts with these different abiotic components. I liked the picture of the horse.

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  4. This paper is well written and a good read. I think an important concept to take away from this paper is climate space, which "quantifies the physical dimensions of fundamental nice" (p. 450). In “Concluding Remarks”, Hutchinson defined and gave examples of a fundamental niche, but Porter and Gates were among the first to quantifying influential parameters of an animal’s fundamental niche.

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  5. The dimensions of radiation, wind, air temperature and humidity, and climate space in general, seem more empirical than the theoretical space discussed in the Bray & Curtis paper. Nice approach! While at first I thought it was silly, the concentric cylinder model does appear to be a good approximation for heat transfer in homeothermic animals. Those little legs and heads sticking out couldn't make that much difference, could they?

    Regarding Porter & Gates' statement, "any organism must be in thermodynamic equilibrium when averaged over a reasonable length of time in order to survive," I find it wonderful that during shorter periods of time, values can swing way up or down, within limits, without resulting in death.

    According to googlescholar.com, this article has been cited 595 times, 31 of these since 2014. In 2014, John Hammond of the UNM Department of Biology was among the authors who cited Porter & Gates in this article: Swimming with Predators and Pesticides: How Environmental Stressors Affect the Thermal Physiology of Tadpoles, http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0098265#references. The reference was in the context of animals' sensitivity to climate change.

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  6. This is really cool stuff - basically using physics and physiological measurements to quantify the climate niches of animals. It brings me back to an environmental biophysics course I took in which we used an equation similar to Porter & Gates' equation 1 to estimate how much food a T. rex would have to eat in order to survive a winter in Wisconsin. I did struggle a bit to interpret the climate diagrams (Figs 13-24) - a lot going in; maybe we can go over in class. I also agree they didn't seem to focus much on water - their focus was more on energy. Maybe this is the result of our dry Southwestern perspective compared to P & G's humid Midwestern perspective.

    This sort of approach seems like it could be used to examine all sorts of interesting questions, some of which P & G touch on e.g. explaining animal behavior and movement patterns, species distribution and habitat modeling, considering trade-offs and implications of body size, color, metabolic rate etc. Body shape also seems like it could be important. They assume animals are cylinders, but the jack rabbit example made me think - are their big ears at least partly a way of shedding excess heat? I also wonder if you could use this to figure out when it's advantageous to be an endotherm v. ectotherm.

    Incidentally, I saw a talk by Warren Porter a few years ago in which he discussed some of this stuff. He's still doing similar things - making models of animals and measuring their thermodynamics. He used his approach to figure out the potential distribution of sea turtles, and also to do some mechanistic species distribution modeling of mosquitoes (or some pathogen carrying insect) in pools of stagnant water in Australia (in watering cans, ran barrels etc). He talked about taking classes, as faculty, in the engineering department to better understand fluid dynamics. Neat guy.

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  7. Like Eric and Noah stated, this paper really grabs my attention in the fact that they are using mathematical and physical models to understand the key variables and then using "climate space" to locate ranges where individuals can survive and thrive. This is the niche space we have been talking about for so long, coined by Hutchinson? Were gates and porter inspired by hutchinson, or did they just happen to think of the same idea but in a different way? Gates and Porter write very in depth at times, but this lack of brevity is probably needed to understand something as complicated as the n-dimensional hypervolume.

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  8. This is definitely an interesting paper, I can see that this is among the first to quantitatively connect the hypervolume with physiology. I thought it a bit odd that they chose the desert iguana (Dipsosaurus dorsalis) as their only poikilotherm example. Dipsosaurus has the highest recorded body temperature of any reptile, they also eat a lot of creosote. They are an oddity among reptiles, especially in North America. Then again, the oddity of Dipsosaurus is probably the reason there was already so much literature available at the time the authors were writing this.

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  9. This paper expand the practical use of the niche concept under the hyper volume definition. The authors did a good job making a quantitative approach (mathematical and physical model) to understand natural constraints that are affecting the realize niche of the species. I think it is a excellent follow-up of the contributions made by Hutchinson and his concept of niche.

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