Speculative biology ramble about things I understand very poorly aldnwkdbskd
Since I only have to understand evo-devo enough to do my speculative biology project, I'm not doing the most thorough job understanding everything. If I figure it out enough to design my own creature, I move on! Otherwise it would take me billions of years to do this. So! This ramble is not necessarily correct about anything, and is entirely in the context of speculative biology rather than a genuine study of earth life!
That being said! Time to ramble! (I'm excited to reread this later and see how much I got wrong, because it'll feel very nice to be able to recognize my mistakes later when I've learned even more!)
It seems that segmentation is, indeed, not a simple thing. I was hoping that each segment would be the result of a discreet set of genes, and that additional segments were the result of those discreet genes duplicating. A gene says "make a segment with kidneys, veins, nerves, a gut tube, etc," and if that gene (or set of genes) gets duplicated, then another segment is added. With that assumption, I figured there'd be at least 3 genes for segments--a "first segment," a "last segment," and a "middle segment" that gets duplicated. I wanted it to work like that, and figured it would, since replicating a head or tail segment would create a weird mouth-mouth or anus-anus. Honestly I was about to give in to that idea, pondering if any of the sphincters of the digestive tract were actually the result of additional head or butt segments being added!
But! Alas! It seems far more complicated than that.
It seems that the segmentation likely arose initially without any discreet segment genes. Though I did read a paper recently about a mulberry moth with a mutation that caused supernumery posterior segments, I think such a mutation is only possible after initial segmentation has been established. I think segmentation originated not as a duplication of an established functional segment, but rather as a division of a non-segmented body. If you knockout a hox gene, it doesn't change the segment count, just the identity of the segments. So segment identity isn't necessarily connected to the creation of the segment. Identity-bestowing genes didn't necessarily arise alongside segment-bestowing genes. The segments--and their identities--could have arisen first, which would need to occur in order to necessitate hox genes...I think.
Anyways, there's another reason I think this. First I'll actually argue against my own point because it's important to get the nuanced picture. (Which is an interesting thing to say when I'm not sure about any of this, only interested in accuracy as long as it serves my ulterior motives of designing aliens wkdjwkdj but I digress.) Sea anemones are considered radially symmetric, and they sure look that way from the outside. But internally they actually seem to have an axis along which different hox genes are expressed. You might be tempted to call this an anterior-posterior axis, which is the only real axis discussed when speaking of hox genes, but that's not really right. Not only do sea anemones have a blind-gut, and thus an oral-aboral axis rather than an anterior-posterior axis--but the linear expression of hox genes doesn't even occur along the oral-aboral axis! To my great surprise they express across the organism horizontally. The only way I can orient this in comparison to a through-gut animal is to wonder if the through-gut evolved not from a continuation of the blind-gut through a secondly formed orifice, but rather from a re-fusing of the middle section of the oral opening of the sea anemone.
I think this is especially possible considering that the planula larva stage of at least one sea anemone experiences a complete fusion and closing of the blastopore. Surely it would be possible for it to only fuse partially, in the middle, leaving two holes, at the ends of the hox gene expression axis. The trouble is that I have no idea how the hox genes express during embryogenesis in sea anemones. Additionally, if the through-gut formed as a central partial fusion of a blind-gut, I'd expect to see evidence of that in the blastopore formation of animals with an alimentary canal. I'd expect a single blastopore to form and then fuse in the center, and then the two pores to migrate to opposite sides. I...don't think that's how alimentary canals are formed? But honestly I've not looked into gastrulation of "higher" animals much, since I am not making those in my world yet.
Regardless, I've noticed that the gastrulation of even "primitive" cnidarians is extremely complex and implies a fascinating evolutionary history. Of course, one must be careful not to fall into the trap of the "theory" of recapitulation, especially since it is tempting to assume that all the evolutionary history could be stored in the process of embryogenesis. But! That is not true. A great deal is obscured through efficient losses and obfuscating mutations and adaptations. All that being said, it is interesting that the blastopore fuses. Why not just have the single layer duplicate and then have the inner layer undergo determination to become the inner layer? Why use invagination and migration? Surely it is a clue to a possible ancestor that experienced the migration and invagination without the fusion. And, perhaps. Either way, it makes me really appreciate the complexity of an animal considered very primitive. Not only are the cnidocytes incredible proof of a complex life form, but even the embryogenesis is convoluted and meandering in its goal, indicative of a long evolutionary history.
Meanwhile, sponges. Sponges seem to have a vast number of strategies for embryogenesis. They aren't considered to have true tissues, and if you study them, it makes you ask, "Wait, what IS a tissue, then?" I didn't look that up yet. But sponges have their cells organize and differentiate. They form a skin-like layer and an inner digestive layer, and a mesohyl between. This is lovely! I think a mesohyl would be an ideal precursor for the salt-regulation system of kidneys and nephrotic systems and things, or perhaps a sort of primitive circulatory system for oxygen and nutrient distribution and waste collection. It reminds me of the extracellular matrix in my own Mesohydrus clade, named after its own similar inner fluid layer! To think that sponges have something like that is truly wonderful.
Again I wonder about the origin of a larval stage. I asked myself, "Just how basal is a larval stage?" when learning about the planula larvae of cnidarians. It seems that many early multicellular animals could have benefited from a larval stage because many adult stages are sessile on account of not having the necessary support for a large nervous system that would allow movement. It seems that early animals may have had to choose between mobility and size. Size does protect you from predators but it is far from the only incentive. In columnar filter feeders, they use a chimney-shaped feeding tube because if the current is faster at one end it will passively draw water through the tube, thanks to my favorite principle, Bernoulli's principle! So a larger size could allow an organism to reach into a different height of water to allow a greater difference in flow velocity over each orifice. This is why several of my own organisms have convergently evolved to be long tall tubes. Interestingly, one of my tubes formed from a barely-motile meandering mouth not unlike a sea anemone, while another arose from a motile worm-like creature that secondarily lost most of its motility in exchange for a slow metabolism and passive filter feeding.
I find myself wondering how basal larvae are, because the easiest way for a sessile animal to spread is by having a small larval stage that is still small enough for coordinated movement, often able to survive on nutrients from the parent while it seeks new habitat. Then it can land somewhere new and afford to develop into an adult and propagate. This is especially curious in placozoans which reproduce through strange means I do not understand. They seem to have a perplexing combination of traits that only make sense to me if they are the result of losses of some traits. They have distinct tissue layers, but no dedicated gastric cavity. Are they flat upside-down sea anemones? Is that the origin of their toxins? Why would they lose a dedicated gastric cavity? They form temporary gastric cavities. Is it simply a matter of being flat? Why? How do they develop even? What does placozoan gastrulation look like? Did their ancestors look like sea anemones, or did the ancestor of sea anemones look like placozoans, or did their ancestor look like something between the two, or different altogether? Why haven't they evolved more? I can think of a hundred ways that placozoans could evolve into new niches. Why aren't they any larger? Why haven't some taken on a sessile life? Why haven't they evolved any form of shell? Or more advanced sensory organs? Of course, if it ain't broke, don't fix it--but also, if you made room for placozoans to evolve into new niches, I think they'd do very well. Why haven't they? What prevents them from having even an iota of visible speciation? We've known about placozoans since the 1800s but struggled to even decide that there is more than one species. Why haven't any of them adapted to the different environments they inhabit?
I think if an animal only reproduced through budding, rather than having embryogenesis, it would struggle to evolve. Of course it struggles due to the lessened genetic diversity. But I think much of evolution--especially regarding body planning--requires embryogenesis. I suppose if the animal had a later metamorphosis, then budding would be less limited, but... There needs to be a phase of fluid cell identity, so that mutations can reassign identity to new areas and develop new body plans. The specific locations of limbs and organs and the specific shape of the organism and the identities that determine the features of the dermis and teeth and placement of eyes and ears and numbers of segments--I think that the evolution of these traits is heavily reliant on a stage of morphogenesis that you don't necessarily see with budding. If an individual forms from the already-organized tissue of a parent, it cannot change its identity or form or function nearly as much. Embryogenesis suddenly seems like an amazing tool for multicellularity. I see why most animals do not reproduce simply by ripping in half. And some that do, like planarians, have fascinating regenerative abilities, not even bothering with having a nice clean bud.
Cell migration plays a much greater part in this than I ever knew. Of course I learned that the embryo begins development via cell migration, but that stage is overshadowed by the later phases. The fingers do not develop from the migration of cells into the final shape. They form a club and then the middle cells die to form the fingers. That seems way cooler than migration! But the migration is crucial. I think it's also a clue into the origins of multicellularity. Colonial single-celled colonies may involve migration of aggregating cells rather than directional growth of the already-present cells. This is certainly true in some organisms like slime moulds. Are slime moulds worth studying in the endeavor to understand metazoan evolution? Do any choanoflagellates aggregate like this? Unrelated aggregations obviously would benefit from the chance for sexual reproduction, but at the same time, cooperation is limited by competition. I struggle to balance these two forces in my own world. Cohorts often experience competition with others for mating opportunities and resources. This competition is curiously managed by the development of sexes, which allows a strange relationship to form between members of the sexes, where the presence of a member of the opposite sex can cause competition and reduce cooperation--or the opposite may occur. The evolution of a gamete binary is a fascinating one, and has had such a great impact on the course of evolution that I am at a loss for words to properly express how important it was. But how does that affect unrelated colonial organisms? What sort of circumstance would allow for unrelated colony members to cooperate? I think choanoflagellates may be a worthwhile study. I think that coordinated flagellation would draw in a better current than a single choanoflagellate could alone, especially when paired with a tube and Bernoulli's principle. But how would they manage the competition? Could they? Or are complex colonies mostly populated by clones? Of course there are organisms I do not understand at all that form multicellular colonies. They look so strange, I am intimidated every time I try to study them. Maybe they would provide insight.
How would these organisms approach embryogenesis? Again the formation of a more mobile larva seems highly beneficial. How ancestral is the larval stage? If it is basal to most metazoans, does it have an impact on embryogenesis even in animals with direct development? How does embryogenesis differ in these two types of animals, in closely related and distantly related animals? If insects could evolve larvae from ancestors that only had nymphs, and if amphibians have larva-like stages, then perhaps it is easier than I think to develop a larva. Do any mammals have them? Could you consider marsupial offspring to be larval? They lack the classic traits I have been quietly assigning larva: niche partitioning to avoid competition with the parent, or the ability to relocate in a way that the parents cannot. Marsupial "larvae" rely entirely on their parents which is the opposite of benefiting from niche partitioning or relocating. Then again, some insects do the same, like honeybees, which nurture their larvae diligently. If one of the main benefits of a larval stage is niche partitioning, why do honeybees feed their young? Why don't their offspring have a more direct development? If they could at least move and eat on their own, and wouldn't they do better? Then again, perhaps it would take them longer to grow and become full sized bees if they had nymphs stages rather than larval stages. It must be difficult to balance the benefits of each.
Most animals don't seem to easily swap between strategies. And the strategies they end up with seem to greatly suit them. Caterpillars thrive with their niche partitioning even at the cost of mobility and a thicker outer layer. Cicadas thrive even at the cost of developing adult traits early on while lacking crucial ones such as functional wings. Marsupials do fine. Placental mammals do fine. Amphibians and lizards both do fine.
But I can only think of a few times in the tree of life where a species has evolved to swap which strategy it uses. Each has very good reasons, too. Amphibians had to lay eggs in the water, so their offspring benefited from being specialized for that, even as adults developed into terrestrial forms. Though, why not just make tiny frogs that can immediately come to land? I think some frogs actually do that. Why do most not? Is it niche partitioning at work again? The insects with larval stages mostly benefit greatly from the niche partitioning. And marsupials were...experimenting with life birth, I guess? That's a bad excuse. Why didn't their ancestors utilize live birth in the first place? That's something I do not know. I know mammals are one of many branches of the tree of life that use live birth. Some sea anemones do something that could be called live birth. Some sharks do. Some snakes. Of course eggs are good on land because they survive outside of water but...live birth also solves that. Perhaps it was just chance. Or maybe the ability to lay an egg and then mate again or just enjoy life without being pregnant was worth it, even if mammals did return to live birth. Do any of these strategies interact with embryogenesis in significant ways? I suppose live birth allows for more maternal influences such as epigenetic changes based on the environment during pregnancy. Of course this could still happen somewhat in eggs, since eggs do indeed originate in the parent. Perhaps even signalling could evolve to cause these changes. I wonder if that's ever happened before. Do any egg-laying animals influence the continued development of the offspring? Some almost seem pre-adapted to such a thing, such as eggs that develop into sexes based on temperature. I wonder if any animals incubate their eggs differently based on the current ratio of sexes in the population. Or if one sex is better at dispersal, maybe the parent would influence the development to favor that sex when the environment is stressful, versus favoring the other when the environment is plentiful.
If embryogenesis is such a key factor in evolution of body plans, do any maternal influences cause increased or decreased mutation of related genes? That would be cool!
To tie it back together nearly, one more benefit of larval stages is neoteny. Of course, direct development can also utilize this, but it causes more drastic changes in animals with larval stages, of course. The niche partitioning is almost like a backup, where the parental niche can be entirely abandoned if needed. Are there any tadpoles that never become frogs? Any grubs that never become six-legged adults? And of course, did any planula larvae take over the reproductive stage, utilizing their increased mobility, creating the first fish-like organism from life that was previously sessile in the adult form?