Nereid said:
In short, the endosymbiosis which resulted in
mitochondria is unique in the sense that, as far as we know, the
mitochondrial DNA of all present-day eukaryotes has a single ancestor. This is not true of plastids.
I see. It certainly is not true of a lot of organelles that might be endosymbiotic relics. It is quite conceivably is true of
mitochondria, but I am cautious. For one thing, how are we to distinguish between a unique source being a single event perpetuated throughout
mitochondria-bearing life, and totally independent ancestors independently picking up the same species of eo-
mitochondrial bacteria in different events?
This is not simple-minded niggling. There could be significant implications for the validity of recognition of monophyletic groups.
By way of analogy, suppose that a new strain of E.coli evolved that developed an endosymbiotic relationship with humans, certain birds, and guinea pigs, conferring say, independence from dietary sources of such vitamins as ascorbate and vitamin B 12. Imagine say, in 100 million years a future palaeontologist and comparative physiologist arguing that those groups (or their descendants of course) must surely be monophyletic.
I am not of course, arguing that this is the case, but that we need a great deal of caution. You see, I do have a deep suspicion that in our earliest stages of development of life on this planet, species were not nearly as exclusively separated as we take for granted nowadays. In fact, I am not really speaking about the very earliest days but about the ages leading into early forms or approaches to eukaryotic biology.
Following this line, and keeping within my general scope, the key questions would be (I think) something like: which of the six (YMMV) supergroups of Eukaryota have forms of mitosis which are not, apparently, shared by other supergroups? And, to the extent could we knock the question into a quasi-hypothesis form, to what extent can we say that one form of mitosis is descended from another (vs, say, an independent innovation)?
[...]
If yes to either, then how did the different forms of mitosis evolve?
Here you go beyond what I know or have an inkling of. In fact I am confident that not all we need to know to comment on some of those questions is yet known at all. They are very important questions, not only for palaeontological reasons, but for understanding the very workings of living cells of all kinds.
What I suspect, but here I am groping in such abject ignorance, that I hardly dare dignify it with the term "speculation", is that when some sort of reasonably effective (I am not saying a word about "efficiency") cell division developed in some of our proto-cellular ancestors, that they rapidly developed de facto parapatry. No doubt (well, all right, plenty of doubt, but live with it!) Much of the crucial machinery had by then been developed, in particular microtubules and various other cellular machinery at all near the molecular level.
On those rather generous assumptions it is easy to imagine how independent improvements to cell division could have developed, all of them culminating in one form of mitosis or another, and not necessarily building on each other's developments.
Conversely, if such early cells were able to fuse, or to enter into endosymbiotic relationships, it is quite conceivable that some may have appropriated say, the ciliary or flagellar motors of others, or what ever they used for making centrioles and the like.
Does that sound excessively vaguely inclusive? That is because it is excessively vaguely inclusive out of ignorance. I leave it to you to imagine the depths of comparative molecular biological research still to be addressed before we can achieve anything like confidence in the details.
And I emphasise again that the ignorance I so glibly refer to hear, is not just the current universal ignorance of humanity, but the far more thorough ignorance of my own.
However, what I do like about this proto-speculation, is that to me at least it suggests a picture of how partly independent development of mitotic machinery, differing in details similar to some items that we already have seen about us, might conceivably have evolved. One involving out of another need not have happened. It could well have been parallel development from common ancestors.
The differences might have been due, not simply to differences in efficiency, but differences in circumstances. Mitosis within nuclear membranes for example might well be an adaptation suited to the environment with in a synkaryon. It might have happened many times, using the same machinery. To us it would look persuasively monophyletic.
What I'm interested in is whether meiosis evolved only once, independently. Or is it like the eye, something which evolved independently many, many times?
Obviously I cannot tell you that, as well you realize, but I find it hard to imagine an early world in which some of the assumptions that I make would not apply in some form or degree. In particular, no matter how wrong my assumptions or deficient my imagination, I find it hard to imagine the mechanism or any of its equivalents being single-event adaptations.
Note that this is totally independent of the question of how many of the component mechanisms and resources may have developed once only. For example, you mention the eye. As I understand it, eyes as structures may have developed umpteen times beyond those that we have identified, but it looks persuasively as though the rhodopsin light sensing mechanism or some precursor, may indeed have evolved just once. Similarly, mitosis, even some forms of mitosis that we regard as effectively identical, might indeed be highly polyphyletic, but all using microtubules of monophyletic origin.
I hadn't thought to ask about mitosis, partly because I'd assumed (wrongly?) that we have no way whatsoever of understanding its origin (today anyway), but with meiosis we may have some possible approaches (e.g. the genetic machinery which is involved).
Well yes, but I don't see why say, the genetic machinery should be dramatically more diagnostic, or at least more suggestive, than comparative work on the machinery of cytokinesis, spindle formation and the like.
Having speculated as far as I have, I do not expect to see resolution of many such questions in my lifetime, but maybe in the lifetimes of my sons. I certainly have been very surprised by some successes that I never had expected, and perhaps more surprised by the routes by which they were discovered.
No doubt ... but there's no way to know, is there?
Not yet!
This all started, for me, when reflecting on the uniqueness of life of Earth, Sagan (or Ward) vs Mayr (or Gould), if you will.
Right. This takes me back to some rather vigorous debates I have engaged in, concerning panspermia.
Something to think about!

Cheers,
Jon