Forever Young

How Evolution Made Baby-faced Humans & Adorable Dogs

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Who among us hasn’t looked at the big round eyes of a child or a puppy gazing up at us and wished that they’d always stay young and cute like that? You might be surprised to know that this wish has already been partially granted. Both you as an adult and your full-grown dog are examples of what’s referred to in developmental biology as paedomorphosis (“pee-doh-mor-fo-sis”), or the retention of juvenile traits into adulthood. Compared to closely related and ancestral species, both humans and dogs look a bit like overgrown children. There are a number of interesting reasons this can happen. Let’s start with dogs.

When dogs were domesticated, humans began to breed them with an eye to minimizing the aggression that naturally exists in wolves. Dogs that retained the puppy-like quality of being unaggressive and playful were preferentially bred. This caused certain other traits associated with juvenile wolves to appear, including shorter snouts, wider heads, bigger eyes, floppy ears, and tail wagging. (For anyone who’s interested in a technical explanation of how traits can be linked like this, here’s a primer on linkage disequilibrium from Discover. It’s a slightly tricky, but very interesting concept.) All of these are seen in young wolves, but disappear as the animal matures. Domesticated dogs, however, will retain these characteristics throughout their lives. What began as a mere by-product of wanting non-aggressive dogs has now been reinforced for its own sake, however. We love dogs that look cute and puppy-like, and are now breeding for that very trait, which can cause it to be carried to extremes, as in breeds such as the Cavalier King Charles spaniel, leading to breed-wide health problems.

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An undeniably cute Cavalier King Charles spaniel, bred for your enjoyment. (Via Wikimedia Commons)

Foxes, another type of wild dog, have been experimentally domesticated by scientists interested in the genetics of domestication. Here, too, as the foxes are bred over numerous generations to be friendlier and less aggressive, individuals with floppy ears and wagging tails – traits not usually seen in adult foxes – are beginning to appear.

But I mentioned this happening in humans, too, didn’t I? Well, similarly to how dogs resemble juvenile versions of their closest wild relative, humans bear a certain resemblance to juvenile chimpanzees. Like young apes, we possess flat faces with small jaws, sparse body hair, and relatively short arms. Scientists aren’t entirely sure what caused paedomorphosis in humans, but there are a couple of interesting theories. One is that, because our brains are best able to learn new skills prior to maturity (you can’t teach an old ape new tricks, I guess), delayed maturity, and the suite of traits that come with it, allowed greater learning and was therefore favoured by evolution. Another possibility has to do with the fact that juvenile traits – the same ones that make babies seem so cute and cuddly – have been shown to elicit more helping behaviour from others. So the more subtly “baby-like” a person looks, the more help and altruistic behaviour they’re likely to get from those around them. Since this kind of help can contribute to survival, it became selected for.

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You and your dog, essentially. (Via The Chive)

Of course, dogs and humans aren’t the only animals to exhibit paedomorphosis. In nature, the phenomenon is usually linked to the availability of food or other resources. Interestingly, both abundance and scarcity can be the cause. Aphids, for example, are a small insect that sucks sap out of plants as a food source. Under competitive conditions in which food is scarce, the insects possess wings and are able to travel in search of new food sources. When food is abundant, however, travel is unnecessary and wingless young are produced which grow into adulthood still resembling juveniles. Paedomorphosis is here induced by abundant food. Conversely, in some salamanders, it is brought on by a lack of food. Northwestern salamanders are typically amphibious as juveniles and terrestrial as adults, having lost their gills. In high elevations where the climate is cooler and a meal is harder to come by, many of these salamanders remain amphibious, keeping their gills throughout their lives because aquatic environments represent a greater chance for survival. In one salamander species, the axolotl (which we’ve discussed on this blog before), metamorphosis has been lost completely, leaving them fully aquatic and looking more like weird leggy fish than true salamanders.

axolotl_ganz
An axolotl living the young life. (Via Wikimedia Commons)

So paedomorphosis, this strange phenomenon of retaining juvenile traits into adulthood, can be induced by a variety of factors, but it’s a nice demonstration of the plasticity of developmental programs in living creatures. Maturation isn’t always a simple trip from point A to point B in a set amount of time. There are many, many genes at play, and if nature can tweak some of them for a better outcome, evolution will ensure that the change sticks around.

Sources

*Header image by: Ephert – Own work, CC BY-SA 4.0, https://commons.wikimedia.org/w/index.php?curid=39752841

An Inconvenient Hagfish

1280px-eptatretus_stoutii

We think of scientific progress as working like building blocks constantly being added to a growing structure, but sometimes a scientific discovery can actually lead us to realize that we know less than we thought we did. Take vision, for instance. Vertebrates (animals with backbones) have complex, highly-developed “camera” eyes, which include a lens and an image-forming retina, while our invertebrate evolutionary ancestors had only eye spots, which are comparatively very simple and can only sense changes in light level.

At some point between vertebrates and their invertebrate ancestors, primitive patches of light sensitive cells which served only to alert their owners to day/night cycles and perhaps the passing of dangerous shadows, evolved into an incredibly intricate organ capable of forming clear, sharp images; distinguishing minute movements; and detecting minor shifts in light intensity.

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Schematic of how the vertebrate eye is hypothesized to have evolved, by Matticus78

In order for evolutionary biologists to fully understand when and how this massive leap in complexity was made, we need an intermediate stage. Intermediates usually come in the form of transitional fossils; that is, remains of organisms that are early examples of a new lineage, and don’t yet possess all of the features that would later evolve in that group. An intriguing and relatively recent example is Tiktaalik, a creature discovered on Ellesmere Island (Canada) in 2004, which appears to be an ancestor of all terrestrial vertebrates, and which possesses intermediate characteristics between fish and tetrapods (animals with four limbs, the earliest of which still lived in the water), such as wrist joints and primitive lungs. The discovery of this fossil has enabled biologists to see what key innovations allowed vertebrates to move onto land, and to precisely date when it happened.

There are also species which are referred to as “living fossils”, organisms which bear a striking resemblance to their ancient ancestors, and which are believed to have physically changed little since that time. (We’ve actually covered a number of interesting living fossils on this blog, including lungfish, Welwitschia, aardvarks, the platypus, and horseshoe crabs.) In the absence of the right fossil, or in the case of soft body parts that aren’t usually well-preserved in fossils, these species can sometimes answer important questions. While we can’t be certain that an ancient ancestor was similar in every respect to a living fossil, assuming so can be a good starting point until better (and possibly contradictory) evidence comes along.

So where does that leave us with the evolution of eyes? Well, eyes being made of soft tissue, they are rarely well preserved in the fossil record, so this was one case in which looking at a living fossil was both possible and made sense.

Hagfish, which look like a cross between a snake and an eel, sit at the base of the vertebrate family tree (although they are not quite vertebrates themselves), a sort of “proto-vertebrate.” Hagfish are considered to be a living fossil of their ancient, jawless fish ancestors, appearing remarkably similar to those examined from fossils. They also have primitive eyes. Assuming that contemporary hagfishes were representative of their ancient progenitors, this indicated that the first proto-vertebrates did not yet have complex eyes, and gave scientists an earliest possible date for the development of this feature. If proto-vertebrates didn’t have them, but all later, true vertebrates did, then complex eyes were no more than 530 million years old, corresponding to the time of the common ancestor of hagfish and vertebrates. Or so we believed.

hagfish
The hagfish (ancestors) in question.  Taken from: Gabbott et al. (2016) Proc. R. Soc. B. 283: 20161151

This past summer, a new piece of research was published which upended our assumptions. A detailed electron microscope and spectral analysis of fossilized Mayomyzon (the hagfish ancestor) has indicated the presence of pigment-bearing organelles called melanosomes, which are themselves indicative of a retina. Previously, these melanosomes, which appear in the fossil as dark spots, had been interpreted as either microbes or a decay-resistant material such as cartilage.

This new finding suggests that the simple eyes of living hagfish are not a trait passed down unchanged through the ages, but the result of degeneration over time, perhaps due to their no longer being needed for survival (much like the sense of smell in primates). What’s more, science has now lost its anchor point for the beginning of vertebrate-type eyes. If an organism with pigmented cells and a retina existed 530 million years ago, then these structures must have begun to develop significantly earlier, although until a fossil is discovered that shows an intermediate stage between Mayomyzon and primitive invertebrate eyes, we can only speculate as to how much earlier.

This discovery is intriguing because it shows how new evidence can sometimes remove some of those already-placed building blocks of knowledge, and how something as apparently minor as tiny dark spots on a fossil can cause us to have to reevaluate long-held assumptions.

Sources

  • Gabbott et al. (2016) Proc. R. Soc. B. 283: 20161151
  • Lamb et al. (2007) Nature Rev. Neuroscience 8: 960-975

*The image at the top of the page is of Pacific hagfish at 150 m depth, California, Cordell Bank National Marine Sanctuary, taken and placed in the public domain by Linda Snook.

Sex & the Reign of the Red Queen

Tenniel_red_queen_with_alice

“Now, here, you see, it takes all the running you can do to keep in the same place.”

From a simple reproductive perspective, males are not a good investment. With apologies to my Y chromosome-bearing readers, let me explain. Consider for a moment a population of clones. Let’s go with lizards, since this actually occurs in lizards. So we have our population of lizard clones. They are all female, and are all able to reproduce, leading to twice the potential for creating more individuals as we see in a species that reproduces sexually, in which only 50% of the members can bear young. Males require all the same resources to survive to maturity, but cannot directly produce young. From this viewpoint alone, the population of clones should out-compete a bunch of sexually-reproducing lizards every time. Greater growth potential. What’s more, the clonal lizards can better exploit a well-adapted set of genes (a “genotype”); if one of them is well-suited to survive in its environment, they all are.

Now consider a parasite that preys upon our hypothetical lizards. The parasites themselves have different genotypes, and a given parasite genotype can attack certain host (i.e. lizard) genotypes, like keys that fit certain locks. Over time, they will evolve to be able to attack the most common host genotype, because that results in their best chance of survival. If there’s an abundance of host type A, but not much B or C, then more A-type parasites will succeed in reproducing, and over time, there will be more A-type parasites overall. This is called a selection pressure, in favour of A-type parasites. In a population of clones, however, there is only one genotype, and once the parasites have evolved to specialise in attacking it, the clones have met their match. They are all equally vulnerable.

The sexual species, however, presents a moving target. This is where males become absolutely worth the resources it takes to create and maintain their existence (See? No hard feelings). Each time a sexual species mates, its genes are shuffled and recombined in novel ways. There are both common and rare genotypes in a sexual population. The parasite population will evolve to be able to attack the most common genotype, as they do with the clones, but in this case, it will be a far smaller portion of the total host population. And as soon as that particular genotype starts to die off and become less common, a new genotype, once rare (and now highly successful due to its current resistance to parasites), will fill the vacuum and become the new ‘most common’ genotype. And so on, over generations and generations.

Both species, parasite and host, must constantly evolve simply to maintain the status quo. This is where the Red Queen hypothesis gets its name: in Wonderland, the Red Queen tells Alice, “here, you see, it takes all the running you can do to keep in the same place.” For many years, evolution was thought of as a journey with an endpoint: species would evolve until they were optimally adapted to their environment, and then stay that way until the environment changed in some fashion. If this was the case, however, we would expect that a given species would be less likely to go extinct the longer it had existed, because it would be better and better adapted over time. And yet, the evidence didn’t seem to bear this prediction out. The probability of extinction seemed to stay the same regardless of the species’ age. We now know that this is because the primary driver of evolution isn’t the environment, but competition between species. And that’s a game you can lose at any time.

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Passionflower. Photo by Yone Moreno on Wikimedia Commons.

Now the parasite attacking the lizards was just a (very plausible) hypothetical scenario, but there are many interesting cases of the Red Queen at work in nature. And it’s not all subtly shifting genotypes, either; sometimes it’s a full on arms race. Behold the passionflower. In the time of the dinosaurs, passionflowers developed a mutually beneficial pollinator relationship with longwing butterflies. The flowers got pollinated, the butterflies got nectar. But then, over time, the butterflies began to lay their eggs on the vines’ leaves. Once the eggs hatched, the young would devour the leaves, leaving the plant much the worse for wear. In response, the passionflowers evolved to produce cyanide in their leaves, poisoning the butterfly larvae. The butterflies then turned the situation to their advantage by evolving the ability to not only eat the poisonous leaves, but to sequester the cyanide in their bodies and use it to themselves become poisonous to their predators, such as birds. The plants’ next strategy was to mimic the butterflies’ eggs. Longwing butterflies will not lay their eggs on a leaf which is already holding eggs, so the passionflowers evolved nectar glands of the same size and shape as a butterfly egg. After aeons of this back and forth, the butterflies are currently laying their eggs on the tendrils of the passionflower vines rather than the leaves, and we might expect that passionflowers will next develop tendrils which appear to have butterfly eggs on them. These sorts of endless, millennia-spanning arms races are common in nature. Check out my article on cuckoos for a much more murderous example.

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Egg-like glands at the base of the passionflower leaf (the white dots on my index finger).

Had the passionflowers in this example been a clonal species, they wouldn’t likely have stood a chance. Innovations such as higher-than-average levels of cyanide or slightly more bulbous nectar glands upon which defences can be built come from uncommon genotypes. Uncommon genotypes produced by the shuffling of genes that occurs in every generation in sexual species.

And that, kids, is why sex is such as fantastic innovation. (Right?) Every time an illness goes through your workplace, and everybody seems to get it but you, you’ve probably got the Red Queen (and your uncommon genotype) to thank.

 

Sources

  • Brockhurst et al. (2014) Proc. R. Soc. B 281: 20141382.
  • Lively (2010) Journal of Heredity 101 (supple.): S13-S20 [See this paper for a very interesting full explanation of this links between the Red Queen hypothesis and the story by Lewis Carroll.]
  • Vanderplank, John. “Passion Flowers, 2nd Ed.” Cambridge: MIT Press, 1996.

*The illustration at the top of the page is by Sir John Tenniel for Lewis Carroll’s “Through the Looking Glass,” and is now in the public domain.

Floral Invasion

Onam_Flower_Arrangement            Throughout evolution, there have been, time and time again, key biological innovations that have utterly changed history thereafter. Perhaps the most obvious is the one you’re using to read this; the human brain. The development of the anatomically modern human brain has profoundly changed the face of the planet and allowed humans to colonize nearly every part of the globe. But an equally revolutionary innovation from an earlier time stares us in the face each day and goes largely unremarked upon. Flowers. (Stay with me here, guys… ) We think of them as mere window dressing in our lives. Decorations for the kitchen table. But the advent of the flowering plants, or “angiosperms”, has changed the world profoundly, including allowing those magnificent human brains to evolve in the first place.

 

Angiosperm percentage
From: Crepet & Niklas (2009) Am. J. Bot. 96(1):366-381

Having arisen sometime around the late Jurassic to early Cretaceous era (150-190 million years ago), angiosperms come in every form from delicate little herbs to vines and shrubs, to towering rainforest canopy trees. They exist on every continent, including Antarctica, which even humans have failed to develop permanent homes on, and in every type of climate and habitat. They exploded from obscurity to the dominant form of plant life on Earth so fast that Darwin himself called their evolution an “abominable mystery”, and biologists to this day are unable to nail down exactly why they’ve been so incredibly successful. Nearly 90% of all terrestrial plant species alive today are angiosperms. If we measure success by the number of species that exist in a given group, there are two routes by which it can be improved- by increasing the number of distinct species (“speciation”), or by decreasing the rate at which those species go extinct. Let’s take a look at a couple of the features of flowers that have likely made the biggest difference to those metrics.

Picture a world without flowers. The early forests are a sea of green, dominated by ferns, seed ferns, and especially, gymnosperms (that is, conifers and other related groups). Before the angiosperms, reproduction in plants was a game of chance. Accomplished almost exclusively by wind or water, fertilization was haphazard and required large energy inputs to produce huge amounts of spores or pollen grains in order that relatively few would make their way to the desired destination. It was both slow and inefficient.

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The world before flowers. By Gerhard Boeggemann on Wikimedia Commons

The appearance of flowers drew animals into the plant reproduction game as carriers for pollen – not for the first time, as a small number of gymnosperms are known to be insect pollinated – but at a level of control and specificity never before seen. Angiosperms have recruited ants, bees, wasps, butterflies, moths, flies, beetles, birds, and even small mammals such as bats and lemurs to do their business for them. The stunning variety of shapes, sizes, colours, and odours of flowers in the world today have arisen to seduce and retain this range of pollinators. Some plant species are generalists, while others have evolved to attract a single pollinator species, as in the case of bee orchids, or plants using buzz pollination, in which a bumblebee must vibrate the pollen loose with its flight muscles. In return, of course, the pollinators are rewarded with nectar or nutritious excess pollen. Or are at least tricked into thinking they will be. Angiosperms are paying animals to do their reproductive work for them, and thanks to incentivisation, the animals are doing so with gusto. Having a corps of workers whose survival is linked to their successful pollination has allowed the flowering plants to breed and expand their populations and territory quickly, like the invading force they are, and has lowered extinction rates in this group well below that of their competitors. But what happens when you expand into new territory to find that your pollinators don’t exist there? Or members of your own species are simply too few and far between for effective breeding?

Selfing morphology
On the left, a typical outbreeding flower. On the right, a selfing flower of a closely related species. From: Sicard & Lenhard (2011) Annals of Botany 107:1433-1443

Another unique feature that came with flowers is the ability to self-fertilise. “Selfing”, as it’s called, is a boon to the survival of plants in areas where pollinators can be hard to come by, such as very high latitudes or elevations; pollen simply fertilises its own flower or another flower on the same plant. Selfing can also aid sparse populations of plants that are moving into new territories, since another of its species doesn’t need to be nearby for reproductive success. It even saves on energy, since the flower doesn’t have to produce pleasant odours or nectar rewards to attract pollinators. Around half of all angiosperms can self-fertilise, although only 10-15% do so as their primary means of reproduction. Why, you may ask, since it’s such an effective strategy? Well, it’s an effective short term strategy. Because the same genetic material keeps getting reused, essentially, in each successive generation (it is inbreeding, after all), over time the diversity in a population goes down, and harmful mutations creep in that can’t be purged via the genetic mix-and-match that goes on in normal sexual reproduction. Selfing as a sole means of procreation is a slow ticket to extinction, which is why most plants that do it use a dual strategy of outbreeding when possible and inbreeding when necessary. As a short term strategy, however, it can allow a group of new colonists to an area to survive long enough to build up a breeding population and, in cases where that population stays isolated from the original group, eventually develop into a new species of its own. This is how angiosperms got to be practically everywhere… they move into new areas and use special means to survive there until they can turn into something new. I’m greatly simplifying here, of course, and there are additional mechanisms at play, but this starts to give an idea of what an unstoppable force our pretty dinnertable centrepieces really are.

Angiosperms are, above all, adaptable. Their history of utilising all possible avenues to ensure reproductive success is unparalleled. As I mentioned, we have the humble flower to thank for our own existence. Angiosperms are the foundation of the human – and most mammal – diets. Both humans and their livestock are nourished primarily on grasses (wheat, rice, corn, etc.), one of the latest-evolving groups of angiosperms (with tiny, plain flowers that you barely notice and which, just to complicate the point I’m trying to make here, are wind-pollinated). Not to mention that every fruit, and nearly every other type of plant matter you’ve ever eaten also come from angiosperms. They are everywhere. So the next time you buy flowers for that special someone, spare a moment to appreciate this world-changing sexual revolution in the palm of your hand.

Sources

  • Armbruster (2014) AoB Plants 6: plu003
  • Chanderbali et al. (2016) Genetics 202: 1255-1265
  • Crepet & Niklas (2009) American Journal of Botany 96(1): 366-381
  • Endress (2011) Annals of Botany 107: 1465-1489
  • Sicard & Lenhard (2011) Annals of Botany 107: 1433-1443
  • Wright et al. (2013) Proc. Biol. Sci. 280(1760): 20130133

**Top image by Madhutvin on Wikimedia Commons **

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Photo by Bernard Dupont on Wikipedia

Theft: Better Than Sex (Bdelloid Rotifers)

(Via: Wikimedia Commons, Image by: Diego Fontaneto)

Common Name: Bdelloid Rotifers

A.K.A.: Families of Order Bdelloida

Vital Stats:

  • Around 360 asexual species
  • All species likely descended from the same ancestor
  • Common ancestor lived 50-100 million years ago

Found: Fresh water bodies of any size, on every continent, including Antarctica

It Does What?!

Here’s a creature that truly exhibits questionable evolution- as in, the kind that tends to make you go extinct in a hurry. Bdelloid rotifers (the ‘B’ is silent) are microscopic animals found in all kinds of moist, freshwater habitats- puddles, ponds, mossy areas; you name it, they’re probably there. What’s so unusual about these guys is that they’re entirely asexual, and have been for a very, very long time. In fact, bdelloid rotifers are all female, a consequence of how they reproduce.

Don’t drink pond water.
(Via: TopNews.in)

Now, asexual reproduction isn’t so uncommon. If you look at a field of dandelions, chances are, they’re all clones derived from asexual reproduction in a single common ancestor- no second parent needed. Even such advanced creatures as komodo dragons do this periodically- a baby dragon is formed from an unfertilized egg inside the mother. What differentiates bdelloid rotifers from other asexual reproducers is that it’s all they’ve done for the last 50 million years or more. Outside of our friends the rotifers, a species must either have sex from time to time, or face extinction.

Why? Because sex solves two major problems in life (your individual results may vary..). First, it weeds out errors which tend to accumulate in DNA over time. Unlike asexuals, which pass on a copy of a copy of a copy (etc.) of their genes, sperm and egg cells contain DNA which has been mixed and matched via a process called meiosis. The gist of this is that an organism can procreate without necessarily passing on any genetic errors it may have to the next generation. Second, this same process of mixing and matching creates new combinations of DNA sequences, which in turn create the natural variation between individuals that evolution can select for or against.

Not the most visually interesting creatures, these rotifers…
(Via: Natural History Museum)

For example, a genetic combination which caused a polar bear to be born with a white nose would be selected for, since it would make a more effective camouflage for hunting. On the other hand, a combination which gave polar bears big black patches on their fur would be selected against, because they’d have a harder time hunting and would therefore starve more often. Asexuals, however, can neither quickly generate useful new combinations, nor purge their populations of harmful mutations.

So on the surface, it comes as a surprise to biologists that bdelloid rotifers have been able to survive for such an epic amount of time with no sex (in addition to the absence of males, genetic tests are able to show that meiosis hasn’t occurred). However, the rotifers have two impressive ways of dealing with this. First, when times get tough, they already have a pretty good defence mechanism worked out- they just dry up. The rotifer dehydrates itself and forms a dormant cyst in which it can remain in this state until conditions improve. This is called anhydrobiosis.

…but what do you expect from sexless pond scum?
(Via: SpaceTravel.com)

Second, and more importantly, they steal genes. This is the true secret to the successful asexual lifestyle. When a rotifer emerges from dormancy and needs to patch itself up, it’s actually able to incorporate random genetic material from its environment into its own genome. A nearby bacterium, some fungus, a passing bit of rotting leaf? All fair game, apparently. Researchers have found genes from each of these three groups in the rotifer genome. Incorporating these new bits of sequence seems to give rotifers the variation they need to develop new traits and stay off the evolutionary chopping block. In fact, given the success of the bdelloid rotifers – they’ve evolved into over 300 species since giving up sex – and the ease of asexual procreation – no need to find a partner – an argument could be made that when it comes to new genes, theft really is better than sex.

Says Who?

  • Gladyshev et al. (2008) Science 320(5880): 1210-1213
  • Harvard Magazine, Nov.-Dec. 2000 “An Evolutionary Scandal
  • Welch & Meselson (2000) Science 288(5469): 1211-1215
  • Wilson & Sherman (2010) Science 327(5965): 574-576