From the Literature: Oxygen, Temperature, and Giant Insects

I hope everyone liked the giant insects post last week!  It was one of my favorites to write.  The topic is just so fun!  I continue with the subject this week by describing a scientific paper that was released in July.  It combines several things I love (giant insects, aquatic insects, and respiration)  into one manuscript of pure science fabulousness!  Let’s get to it, shall we?

You probably learned as a kid that insects are ectothermic (aka, cold-blooded).  Ectothermic organisms are largely unable to regulate their body temperatures, so their bodies remain close to the temperature of their environments.  As the temperature increases, processes like metabolic rates speed up.  The opposite happens at cold temperatures and everything slows way down.  Ectotherms survive best under a range of temperatures where their body processes work efficiently, but the animal is still able to get everything it needs (food, water, oxygen, etc) from the environment.  They’re like Goldilocks: they don’t like things too hot or too cold and prefer for things to be just right.

Oxygen plays a big role in the interaction of ectotherms with their environments, especially at extreme temperatures.  Let’s consider a hypothetical insect, say a grasshopper.  As the grasshopper gets warmer, its metabolic rate increases and its body processes become more efficient.  However, as the grasshopper’s metabolic rate increases, so does its oxygen consumption.  At some point, the oxygen demand of the grasshopper may become greater than its oxygen availability and all sorts of bad things start to happen as its body processes start to break down.  Oxygen plays a role at very cold temperatures as well, leading scientists to propose that oxygen can set thermal limits (the maximum and minimum temperatures our grasshopper can survive) in ectotherms.

One problem though: terrestrial insects don’t fit the pattern observed in many other ectothermic animals.  This may be because their respiratory systems do not depend on lungs and blood to deliver oxygen to their cells and instead deliver oxygen directly to their cells via a series of tubes that connect to the outer environment.  This creates a terribly efficient system that provides enough oxygen even at high temperatures for many terrestrial insects.  Quite simply, their respiratory system provides enough oxygen even under the worst conditions.  But what if the insects live in oxygen-limited environments, such as water?  Might oxygen play a role in setting those upper thermal limits then?


Image from bilder/030416_dinocras_cephalotes.htm.

Researchers Wilco Verberk and David Bilton considered this question and determined that if the thermal limits of any insects were to be limited by oxygen levels, aquatic insects were the most likely suspects.  So, they chose an insect that requires a low temperature and flowing water as their subject, the stonefly Dinocras cephalotes.  If the maximum temperature the stonefly could tolerate was limited by oxygen consumption, the maximum tolerable temperature would decrease in low oxygen water while it would increase in high oxygen water.  They then developed a simple experiment to determine whether this was the case.

The team placed stoneflies in flow-through chambers in a water tank and ran 10°C water containing various mixtures of oxygen and nitrogen (20% O2/80% N2 = normal, 5% O2 /95% N2 = low oxygen treatment, and 60% O2/40% N2 = high oxygen treatment) through them.  After letting the stoneflies acclimate for an hour, they ramped the temperature of the water up 0.25 degrees per minute until the critical temperature was reached, i.e., the stoneflies started showing signs of thermal stress such as lack of movement and leg twitching.  Then they compared the critical temperatures for each treatment to determine if their hypothesis was correct.

And it was!  They discovered that the upper thermal limit increased almost 3°C in the high oxygen water compared to water containing normal levels of oxygen.  Conversely, the upper thermal limit decreased in low oxygen water by about 1.5°C compared to that at normal oxygen levels.  The conclusion: oxygen levels can set upper thermal limits in larval aquatic insects!

Now you might be wondering why this is exciting or what any of this has to do with giant insects.  The results are interesting for several reasons, but largely because they show that some insects do experience oxygen-induced changes in their upper thermal limits.  This means that, while terrestrial insects might be able to obtain enough oxygen at any temperature to meet their needs, aquatic insects and other insects that live in oxygen limited environments can reach a temperature at which their oxygen demand outstrips the oxygen available to them.  Consider how an insect such as a stonefly gets the oxygen it uses.  They don’t have any spiracles (the pores through which terrestrial insects “breathe”), so oxygen is simply absorbed through the exoskeleton.  Many stoneflies have gills to make this process more efficient (the bigger your body surface, the more oxygen you can absorb from the water), but it’s still a very slow process.  The size of these insects may be limited as a result.  Aquatic insects that rely on absorbing oxygen from the water rather than going to the surface to breathe are also unable to regulate their oxygen uptake very well.  They can do various behaviors to increase the flow of oxygen into their bodies when they become oxygen stressed, but oxygen becomes toxic at very high concentrations.  Aquatic insects can’t do much to prevent oxygen from flowing into their bodies, so this can be a problem.

And this brings us to the giant insect part of the paper.  Verberk and Bilton propose that oxygen limitation at temperature extremes may have contributed to the rise of insect gigantism in the late Carboniferous and early Permian.  This makes sense considering how many of the giant insects were insects that probably had aquatic nymphs (proto-dragonflies, mayflies, and stoneflies, among other aquatic organisms).  The high levels of oxygen at the end of the Palaeozoic meant that oxygen could be absorbed more efficiently by aquatic insects and allowed them to become larger.  I covered this hypothesis last week, so check that post for more details.

Alternatively, Verberk and Bilton suggest that oxygen toxicity may have played a significant role in promoting insect gigantism.  How can an aquatic insect cope with increasing levels of oxygen in water and prevent oxygen poisoning?  They can get bigger!  If insects increased in size as oxygen levels in water rose, then they could counteract the negative effects of high oxygen levels on their bodies.  Oxygen levels at the end of the Palaeozoic were so high that aquatic insects likely had to become very large to prevent oxygen poisoning.  Giant immatures then led to giant adults.  Hence, giant insects that resulted from the limits of their respiratory systems in very high oxygen environments!  It’s a very interesting, new idea.  I suspect many people will do further tests in the future to determine whether this might really have been possible, so we’ll see if it holds up to further study.

I love this hypothesis!  Still, I have to point out that there is one major assumption that the entire hypothesis is built upon, that the giant proto-dragonflies, mayflies, stoneflies, etc had aquatic nymphs.  Modern dragonflies, mayflies, and stoneflies have aquatic immatures, so it’s likely that their predecessors did too.  However, there is no fossil evidence of aquatic nymphs for these groups at the time of the giant insects.  For all we know, the griffenflies and giant mayflies may have had terrestrial nymphs, which would make Verberk and Bilton’s hypothesis fall apart completely.  While the authors did acknowledge this assumption, I think their position would be strengthened if a fossil of even one aquatic immature could be found from that time period.  Without that piece of evidence, I fear this hypothesis is built upon a shaky foundation, one that might not hold up to scrutiny.

But wow!  A new explanation for how giant insects may have evolved!  And focused on the aquatic stages of insects!  You can see why I’m excited by it.  I can’t wait to see the research generated by this paper in the future.  It’s going to make for some very interesting reading!


Literature Cited:

Verberk WC, & Bilton DT (2011). Can oxygen set thermal limits in an insect and drive gigantism? PloS one, 6 (7) PMID: 21818347

This paper is open access!  Full text available online here:


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5 thoughts on “From the Literature: Oxygen, Temperature, and Giant Insects

  1. Don’t fossils form more readily in aquatic environments? Why would the terrestrial/airborne versions of the insects be more common in the fossil record than their (hypothetical) aquatic larvae?

    I suppose the question to ask would be do insect groups that we know have aquatic larvae have those larvae represented in the fossil record? I knew I should have finished reading that book on insect fossils already. : – )

    • The modern dragonflies, etc do have a fossil record of aquatic nymphs. They’re represented in aquatic environments since around, I want to say, the Jurassic or so (maybe even the Triassic in some groups – I’d have to look it up to be sure). But the really old fossils of proto-odonates and other precursors to the modern aquatic insect fauna have no record of aquatic nymphs. You’re right about fossils forming more readily in aquatic environments, so it seems strange that if the giant precursors of aquatic insects had aquatic nymphs that there would be no record of them. So many of the major insect fossil deposits are situated at the former sites of large bodies of water! This is why I have a hard time completely supporting the aquatic nymphs driving gigantism hypothesis. I think it’s a big assumption that the griffenflies and other giant insects of the late Carboniferous and early Permian were aquatic just because their modern descendants are. I personally think it likely they were aquatic, but without fossil confirmation of this I think the hypothesis that Verberk and Bilton propose is weaker than it might be otherwise. Still, it’s an intriguing idea and will lead to lots of interesting research, so I really love the paper.

  2. Maybe they were aquatic, but not from depositional environments. Say they lived in rivers or something. That would reduce their likelihood of fossilization, wouldn’t it?

    But I suppose you’d still expect dead ones to end up in river mouth depositions and the like.

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