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Monday, August 31, 2015

Do snakes sleep?

Do snakes sleep? Do they dream? These may seem like obvious questions, especially since almost every species of mammal, bird, reptile, amphibian, fish, and invertebrate studied has been found to exhibit some kind of resting phase. But sleep is hard to study in snakes, at least in part because they seem never to close their eyes. Consequently, there is shockingly little research on sleep in snakes. A Google Scholar search for the terms "snake+sleep" returns papers about venomous snakebites to sleeping victims, sleepwalkers dreaming about snakes, and papers by Stanford geophysicist Norman H. Sleep on the geology of the Snake River in Idaho. But, despite the dearth of research, I promise this post won't be too much of a snooze...

Human EEG "brainwaves"
Sleep is a behavior that involves an immobile posture, decreased responsiveness to arousing stimuli such as noise and light, and rapid reversibility (the ability to quickly "wake up", as distinct from hibernation or a comatose state). The physiological criterion most frequently used to define sleep is the slowing down of "brain-waves" on an EEG. An EEG (electroencephalogram) measures electrical activity in your brain, which is caused by your brain cells talking to one another. Brain activity, which happens even during sleep, appears as wavy lines on an EEG recording, hence brain 'waves'. When mammals and birds are sleeping, they exhibit two alternating patterns of EEG activity: 1) slow-wave sleep (SWS, also called synchronized, quiet, or non-REM sleep), which is characterized by high amplitude (75-400 μV), low frequency (0.5-4 Hz) EEG waves, and 2) "paradoxical" sleep (PS, also called desynchronized, active, or REM sleep), which is characterized by low voltage (5-10 μV), high frequency (13-30 Hz) EEG waves that are physiologically more like those in awake animals (hence the name "paradoxical"). In humans and cats, paradoxical sleep is associated with rapid-eye movement (REM, measured by electro-oculography or EOG), complete muscle relaxation (measured by electromyography or EMG), muscle twitching, irregular breathing/heartbeat, and, in humans at least, with dreaming.

Lizards wearing EEG-recording equipment while awake and asleep
From Flanigan 1973
Although sleeping patterns are enormously variable across the animal kingdom, most mammals and birds tested exhibit both SWS and PS, or variations on that theme. In some basal mammals and birds (echidnas, platypus, ostriches), eye movement and relaxed muscle tone are associated with both quiet and active sleep. Periods of rest or quiescence associated with EEG changes similar to those seen in mammalian sleep are clearly present in turtles and in crocodilians, but EEG data suggest that these animals do not exhibit REM sleep. Some experiments have found REM-like sleep in lizards, whereas others have not. In experiments where lizards, turtles, and crocodilians were subjected to continuous arousal for 24-48 hours, they appeared to get tired, sleeping more afterwards and producing more high-voltage EEG spikes. Tortoises given the drug atropine, derived from the mandrake plant and used to produce deep sleep in humans since at least the fourth century B.C.E., also produced more spikes, suggesting that EEG spikes are in fact analogous signs of quiet sleep in reptiles and mammals. Interpreting EEG data is complicated because SWS waves differ between mammals and reptiles, perhaps because reptile and mammal brains differ in structure, particularly with respect to the neocortex, the source of these waves in mammals. Furthermore, some reptiles sometimes seem to exhibit sleep-like brain activity when they are awake, perhaps because ectotherms basically fall asleep when they get cold.

Waking (top) and sleeping (bottom) python EEG
and EMG waves. From Peyrethon & Dusan-Peyrethon 1969
The single study of a snake was done by French comparative sleep researchers J. Peyrethon and D. Dusan-Peyrethon (I could not find their full first names), who also studied sleep in fish, caimans, cats, and mice in the 1960s at the Laboratoire de Médecine Expérimentale in Lyon. They used EEG to monitor the brainwaves of a four-foot African Rock Python (Python sebae) over two days. They reported that sleep-like brain waves were produced almost 16 hours a day, increasing to over 20 hours following feeding, and that these brainwaves corresponded with slower breathing and heart rate, some muscle relaxation, and perhaps a lowered behavioral response threshold. They did not see any evidence for active sleep in the EEG. As far as I can tell, this is the only study ever conducted on sleep in a snake.

Snakes do have circadian rhythms, and many snakes are active only at particular times of day. Racers (Coluber), hog-nosed snakes (Heterodon), patch-nosed snakes (Salvadora), and sipos (Chironius) are strictly diurnal, whereas aptly-named nightsnakes (Hypsiglena), broad-headed snakes (Hoplocephalus), and kraits (Bungarus) are strictly nocturnal. But many snakes do not fit nicely into these categories. Good examples include ratsnakes (Pantherophis) and many vipers, but many other snakes may be active at any time of the day or night, depending on the time of year, so it's hard to predict when or for how long they might be expected to sleep. You often observe snakes exhibiting sleep-like behavior, sitting in one spot for hours, days, or even weeks at a time, like the Puff Adder (Bitis arietans) in the video at left. But the thing is, that snake is actually foraging. A viper might sit motionless for many days, such a long time that if a mammal exhibited that same behavior, we might think it was sick or dead! But in fact this is how many snakes forage for prey, hyper-alert to their immediate surroundings, ready to ambush, strike, and envenomate small animals that stray too close. Do they sleep when they are waiting, or are they awake the entire time? Radio-telemetry studies of bushmasters (Lachesis muta) in the wild suggest that they might have strict cycles of attentiveness, "awesomely alert during darkness and almost as if drugged by day", with relatively abrupt transitions each way. On the other hand, many marine mammals and migratory birds do not seem to sleep for long periods of time without suffering any obvious consequences. When engaged in constant activity, these animals close one eye and sleep one half of their brain at a time. Other animals, including perhaps some lizards, sleep one hemisphere at a time in contexts of high predation risk. Might snakes that use sit-and-wait foraging strategies do something similar?

I photographed this Sonoran Lyresnake (Trimorphodon lambda)
during the day, but it was found at night. Their skinny slit-like
pupils enhance their night vision, making distant
objects sharper by increasing the depth of field,
like using a small aperture on a camera lens.
If lyresnakes sleep, it's probably during the day.
How would a researcher tell if a snake was sleeping? Snakes never close their eyes. Or, more accurately, their eyelids are always closed, but they are covered by clear scales. Either in the wild or in captivity, observations of snakes seeming to "wake up" (implying that they were sleeping) are rare: motionless snakes rarely twitch, and other signs of PS are either normal for snakes (such as irregular breathing/heartbeat) or anatomically impossible (REM). You could imagine a series of experiments where an experimenter used EEG and high-speed infrared videography to record the brainwaves and behavioral responses of snakes to arousing stimuli. What stimuli to use is an open question, since snakes don't necessarily respond to bright lights or loud noises even when they're awake. Because snakes inhabit a primarily chemosensory world, it might be possible to wake one up using a smell. The human experience would suggest that the onset of chemosensory signals is inherently too gradual to really be surprising, but this might or might not be true for snakes. What about the infrared sense of some snakes? Could a bright infrared light wake them up? Can snakes see when they're asleep? What would that even be like? Only further studies will tell for sure.

So here's what we know: snakes probably do sleep, perhaps most of the time, but we don't really know when, for how long, how deeply, or whether or not they have paradoxical sleep, including dreaming. Sleep patterns are probably quite diverse across the >3500 species, of which only one has been examined. Many snakes do yawn, but this has been interpreted either as a means to gather chemical cues or to reposition musculoskeletal elements, in contrast with the hypothesized functions of yawning in humans (possibly regulating brain temperature, causing increases in blood pressure, blood oxygen, and/or heart rate in order to improve motor function and alertness, or as a social cue). Sleep is such a basic element of human biology, so if you ask me, the subject of sleep in snakes, and broader questions about the diversity, evolution, and function of sleep across the animal kingdom, should be keeping researchers awake at night.


Thanks to Kendal Morris for suggesting this question, and to Harry Greene, David Cundall, and Gordon Burghardt for sharing their observations.


Ayala-Guerrero, F., & Huitrón-Reséndiz, S. 1991. Sleep patterns in the lizard Ctenosaura pectinata. Physiology & Behavior 49:1305-1307 <link>

Bauchot, R. 1984. The phylogeny of sleep in vertebrates [birds, reptiles, amphibians, fish]. Annee Biologique (France) 23:367-392 <link>

Brischoux, F., Pizzatto, L., & Shine, R. 2010. Insights into the adaptive significance of vertical pupil shape in snakes. Journal of Evolutionary Biology 23:1878-1885 <link>

Campbell, S. S., & Tobler, I. 1984. Animal sleep: a review of sleep duration across phylogeny. Neuroscience & Biobehavioral Reviews 8:269-300 <link>

De Vera, L., González, J., & Rial, R. V. 1994. Reptilian waking EEG: slow waves, spindles and evoked potentials. Electroencephalography and Clinical Neurophysiology 90:298-303 <link>

Flanigan, W. F. 1973. Sleep and wakefulness in iguanid lizards, Ctenosaura pectinata and Iguana iguana. Brain, Behavior, and Evolution 8:417-436 <link>
Greene, H. W., & Santana, M. 1983. Field studies of hunting behavior by bushmasters. Estudios de campo del comportamiento de caza por parte de las cascabelas mudas. American Zoologist 23:897 <link>.

Hartse, K.M. and A. Rechtschaffen. 1974. Effect of atropine sulfate on the sleep-related EEG spike activity of the tortoise, Geochelone carbonaria. Brain, Behavior, and Evolution 9:81-94 <link>

Libourel, P. A., & Herrel, A. 2015. Sleep in amphibians and reptiles: a review and a preliminary analysis of evolutionary patterns. Biological Reviews <link>

Peyrethon, J., & Dusan-Peyrethon, D. 1969. Etude polygraphique du cycle veille-sommeil chez trois genres de reptiles. CR Soc Biol (Paris) 163:181-186 <not available online>

Rattenborg, N. C. 2006. Do birds sleep in flight? Naturwissenschaften 93: 413-425 <link>

Roe, J. H., Hopkins, W. A., Snodgrass, J. W., & Congdon, J. D. 2004. The influence of circadian rhythms on pre-and post-prandial metabolism in the snake Lamprophis fuliginosus. Comparative Biochemistry and Physiology Part A: Molecular & Integrative Physiology 139:159-168 <link>

Siegel, J. M. 2008. Do all animals sleep? Trends in Neurosciences 31: 208-213 <link>

Siegel, J. M., Manger, P. R., Nienhuis, R., Fahringer, H. M., Shalita, T., & Pettigrew, J. D. 1999. Sleep in the platypus. Neuroscience 91: 391-400 <link>

Tauber, E.S., J. Rojas-Ramirez, and R. Hernandez-Peon. 1968. Electrophysiological and behavioral correlates of wakefulness and sleep in the lizard (Ctenosaura pectinata). Electroencephalography and Clinical Neurophysiology 24:424–443 <link>

Creative Commons License

Life is Short, but Snakes are Long by Andrew M. Durso is licensed under a Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License.


Unknown said...

And the situation with night snakes (Hypsiglena) is complicated as well--they surely do rarely at most move about during the day outside of cover, but available evidence shows they mainly capture prey by day, ambushing diurnal lizards from hiding.

Andrew Durso said...

Good point! Do they ever sleep?

Unknown said...

Studying brain waves is nice and all, but have We become so advanced ("advanced", perhaps) that We have forgotten the base of Science: observation? Also; applying logic, common sense, analytical thought, and intuition?

Id est; serpents sleep, and it is indicative via their pupils 'dropping' as though facing down, and appearing to be in total autopilot mode, looking nothing like when they're moving about, foraging, eating or drinking. When stimulated or simply awakened, the pupils rise to normal-appearing mode, and the serpent slightly lifts its head as though becoming attentive. This is the observation part of it, and I've watched for 24 years.

The logical, common sensed, analytical and intuitive part is simply that every animal sleeps, period. We don't need concrete proof of this, such as via technology. As with humans, rest is just as essential as keeping active and fit, as rain is to sunshine, and as war is to peace. Id est; it is the essential balance of Nature and of Life.

Dreaming? That's a tougher one to discern, and if it can be figured out not just in serpents but all animals, it may distinguish just how differently animals sleep.

On a final note, I found the smell stimuli interesting, as I have had otherwise nocturnal species (exempli gratia, Boiga dendrophila) eat in early-mid afternoon when I knew they were asleep in their caves below. Granted, opening the enclosure could have woken them, and this is where brain wave equipment would be necessary - to see if anything changes from opening to closing the cage, or does nothing change until a certain time passes where it can be almost guaranteed the smell of food woke and brought the animal(s) out?


Andrew Durso said...

Thanks for your thoughtful comment, Tim. I would add that, although in general I agree with you that we shouldn't use technology just because we can and that we should always be making observations and thinking about how they can suggest clever experiments to test hypotheses, we do need technology and physiological experiments, and sometimes, "common sense" and "intuition" can lead us astray. It's certainly been shown that snakes sleep. But, I challenge your assumption that "every animal sleeps, period". For instance, I recently learned about some Antarctic insects that hibernate for most of the year, but don't sleep or have a circadian rhythm during the short Antarctic summer during which they are active 24 hours a day. Also, without technology, we wouldn't know about pelagic birds and mammals that sleep half their brains at a time, seeming behaviorally never to sleep.

Kobelkova, A., S. G. Goto, J. T. Peyton, T. Ikeno, R. E. Lee, and D. L. Denlinger. 2015. Continuous activity and no cycling of clock genes in the Antarctic midge during the polar summer. Journal of Insect Physiology 81:90-96.

Unknown said...

And I thank You for Your thoughtful, insightful response, Andrew. :) Indeed; relying too much on common sense and intuition can become 'cloudy', and it was not my intention to convey the notion that science should be based on common sense and intuition exclusively. Technology definitely has its benefits, not least to show Us physiological aspects that We cannot see (on the outside).

I suppose when I stated 'every animal sleeps, period', I admittedly was not thinking of, and therefore not including insects. It's not that I put insects below vertebrates, I was just simply ignorant LOL (or perhaps too tired). It was probably also--perhaps subconsciously--the connotation of "animal" growing up which drew a line between certain groups/types/kinds of animals, perhaps between vertebrates and insects (not invertebrates in general, just insects); this is obviously wrong, for insects are a part of Kingdom: Animalia.

So; Antarctic insects that hibernate but don't sleep...I still didn't know about them either way, so You got me there! But would it be more like a reptilian brumation then, not hibernation? ...which is not a coma-like sleep like hibernation is. And my use of 'sleep' is actually more broad, incorporating, say, 'autopilot' mode which is not completely unconscious, or in the case of these pelagic birds and mammals: on 'standby' mode so to speak. I should have specified.

Lastly, I'm not surprised about these pelagic birds and mammals sleeping half of their brain at a time; this would explain the Albatross (a pelagic bird) sleeping while flying...rather; the source of that ol' fact I learned years ago probably didn't realize it was only sleeping half of its brain at a time (because it definitely hadn't specified this...it was a sign at the Forest Park Zoo in Springfield, MA)...I sure as heck didn't realize it till Your comment just now, but it makes sense. And yeah, something like pelagic birds flying way above the ocean can't be discerned to be sleeping or not--how can You see and even think of the possibility/ability which would sound absurd to most people initially, until proof tells them--and only technology was able to furnish the proof.

Andrew Durso said...

Tim, it's certainly true that there are a myriad of wonderful exceptions to and twists on every "rule" in nature—my favorite thing about biology is those exceptions! I can be pretty excited about pointing them out. Thanks again for the thoughtful comments.

Unknown said...

And it's good to be so excited and passionate about something! :) Welcome, and thank You for the great articles.

Andrew Durso said...

A good article arguing that the distinction between the terms 'hibernation' and 'brumation' is hazy at best and unnecessarily jargon-y:


Unknown said...

Sure enough, researching the etymology of the two terms: brumation and hibernation reveals no specification/difference between them in regards to the physiological/behavioural nature of reptiles (or any creature for that matter) over Winter. But there is a physiological/behavioural difference between reptiles and mammals in that the former does not undergo a coma-like sleep and will instead be sluggishly active, occasionally drinking and even basking at the opening of their hibernaculum on warm spells.

And this is where semantics must've been put aside for simplicity sake however long ago, to use simple terms in distinguishing a (physiological/behavioural) difference between reptiles and mammals in their overwintering nature irrespective of the literal, etymological meaning of the terms. Perhaps it is similar to old-timers (still) interchangeably using poisonous and venomous for the physiological process of inJECting venom. NOTE: It is important to specify 'physiological process of inJECting' since it has been recently discovered that at least five (5) genera of serpents are poisonous (and coincidentally all five are venomous to some degree, or inject toxins*), in which the physiological process of poisonous is inGESting. But old-timers weren't/aren't referring to toxins sequestered in the nuchal glands when using "poisonous", haha; they were/are referring to serpents' fangs and venom glands.

In conclusion: if not "brumation", the so-called "experts" need to find (or create) a term that identifies and distinguishes reptiles from mammals in their overwintering physiology/behaviour...there is a difference obviously, and I want a word to describe it!

Thank You for the follow-up comment and link, though; I am big into linguistics and quite the etymological prick actually LOL, so I like learning facts like this! Admittedly/shamefully for this 'linguistic connoisseur' who took three (3) of Latin, haha, I don't think I ever rooted out/researched the etymology of either term, at least too long ago for me to recall (and obviously forgot), and therefore never realized brumation technically doesn't distinguish anything, HA!

*Prepare for a digression-of-an-asterisk note completely irrelevant to this topic LOL:

Heterodon platyrhinos at least, if not the entire genus Heterodon is actually not mildly venomous, but rather mildly toxic since the colour of their secretions is clear whereas venom is always some shade of yellow, not to mention significant composition differences. An excerpt of greater elaboration, from Linda Krulikowski's amazing book: Snakes of New England (with the amazing Harvey B. Lillywhite contributing):

"The [Duvernoy's] gland is composed of branched tubules rather than simple groupings of cells, and is situated immediately under the skin, above and near the angle of the jaw. It is derived from the same tissue as tooth enamel. The Duvernoy's gland opens by a duct at the base of the posterior, enlarged teeth. The secretions from the glands flow down the enlarged, rear teeth and into the prey, by the chewing motions of the snake. The secretions are introduced slowly into the victim, by indirect pressure from the nearby jaw-closing muscles (Green. 1997: 78-95). The secretions of Duvernoy's glands are colorless, whereas true venom glands secretions are typically some shade of yellow. These clear secretions immobilize the victim and help the snakes digest the prey."

Unknown said...

three (3) years* of Latin [forgot a word, sorry, and evidently cannot Edit comments...?] :P

Andrew Durso said...

Thank you again Tim for your thoughtful comments.

1. My aim in pointing out that hibernation and brumation are not so different is not to suggest that there are no physiological/behavioral differences between reptiles & mammals. Rather, I wanted to emphasize that reptilian hibernation and mammalian hibernation are not two distinct categories but rather part of an evolutionary continuum of deep-sleep strategies used by vertebrates, one that is intimately tied to the ectothermy-endothermy spectrum. For instance, hibernating echidnas have periods of arousal, during which they often move to other locations. Probably monotreme hibernation is the closest we can get to a living organism with similar physiology to that of the common ancestors of all amniotes.

Regardless of what I or any etymological or biological expert says, "brumation" will continue to be used for herps and "hibernation" for mammals, and there's nothing wrong with that. But simplifying the biological world into mutually-exclusive categories often denies us the true complexity of the underlying reality, and gives the misleading impression that reptilian and mammalian physiology/behavior were independently created in isolation from one another, when in fact they were not.

2. Your point about the similarity between the terms "hibernation/brumation" and "poisonous/venomous" is a good one, and a good example of how term use can change over time. Again I would argue that toxin injection has evolved numerous times and there are grey areas (such as the delivery of skin gland toxins by the pointed ribs of Pleurodeles waltl and other salamandrids). But, again, the distinction is useful and will continue.

3. I don't think that anyone uses the color of snake oral secretions as a useful way to classify them as venom or not venom. I think anyone would agree that the chemical composition and function are more important characteristics than the color. I looked into it and evidently the venom of many vipers is colorless when they are first born, and the venom of some adult Asian cobras is colorless as well. I suspect you'll agree that these colorless secretions are still venom. The differences between the composition of Heterodon venom and that of other snakes are no greater than would be expected given their evolutionary distance from those other snakes. Once more, the distinction between "true venom" and "whatever non-front-fanged colubroid snakes have" is A. only a matter of degree, not mutually-exclusive categories, and B. categorized primarily, if not solely, by the life-threatening potential towards humans (and, again, this distinction is useful; in fact I think that describing snakes that don't pose medically-significant threats to humans as "venomous" is misleading and confusing, which is why I often try to point out that venomous/non-venomous is also an evolutionary continuum).

4. Regarding the sequestration of toxins into nuchal glands, we have conclusive evidence only that Rhabdophis tigrinus sequesters bufadienolides into its nuchal glands. We have every reason to believe that other Rhabdophis, and the two closely-related genera Macropisthodon and Balanophis, also use their nuchal/nucho-dorsal glands for this purpose (although the toxins may vary). Fourth, there is evidence that Thamnophis retain tetrodotoxin in their livers for up to two months. But as far as I know, to date this is the extent of our knowledge of the diversity of non-injected chemical defenses in snakes.

5. Sorry about not being able to edit comments; it's one of the several drawbacks of using the Blogger platform.

Thanks for reading and writing!

Mike Pingleton said...

Seems like an interesting project for someone. I wonder how tongue-flicks figure into this? Can you assume that no flicks for a certain time period = sleep state?


Andrew Durso said...

Definitely. Good idea—that's possible, and testable. Hopefully someday someone will take it on.

Bill said...

Nice summary regarding sleep in snakes, Andrew. Here is a paper that defines what a venom is and distinguishes it from poisons and toxungens: https://tinyurl.com/yexrsd86. - Bill Hayes