Wednesday, February 13, 2013

Just Leave The “Father” Line Blank

Biology concepts – apomixis, automixis, genomic imprinting, haplodiploid, facultative and obligate parthenogenesis

Kids questions can be exasperating, exhilarating,
and problematic –all at the same time. Questions
about biology are especially difficult because you
never know how much information to give at what
age. My advice – give the simplest answer that will
stimulate additional questions. Too much detail can
be a turn off to a kid and can lead you into subjects
that you aren’t ready to tackle with them. If you don’t
know the answer – find it out. Your kids should see
you demonstrate looking for an answer and learning.
“Mommy, why is the sky blue? Daddy, if atoms are mostly empty space, then why are objects solid?” These are questions with which every parent must deal. Unless you are familiar with Raleigh scattering or the quantum structure of the atom, you’re going to have to make something up. Use big words – it will confuse them into moving on to something else, and you get to look like you know something.

Parthenogenesis is a subject that baffles a lot of people for a lot of reasons, mostly because we know little about it yet. What has science’s response been to this lack of knowledge – give everything a new name and bury people in mountains of terminology. Jargon is job security after all.

Last week we saw that parthenogenesis, while an exception, is not as rare as we once thought. Now let’s take some time to get down and dirty and look at the in and outs of abandoning sex. We’ll use examples to keep the vocabulary monster at bay.

The whole point of parthenogenesis is to make an unfertilized egg develop into a whole organism. How can an egg develop on it own? In general, haploid eggs are useless (exception alert!) so moms need to construct a diploid or polyploid egg in order for parthenogenesis to have a chance.

One way is for mom to forego meiosis and produce diploid eggs. The term for this is apomixis (apo = free from, and mixis = mixing). It basically means "with no mixing of chromosomes;" neither by homologous recombination nor by random assortment in meiosis. Therefore, offspring produced by apomictic parthenogenesis are clones of their mothers.

Automixis results in half clones. The mother’s eggs
go through meiosis, so the joining of different
products to regain diploidy will necessarily join
unlike chromosomes. Therefore, the offspring can’t
be a full clone of the mother. The mixing can come
from joining two eggs that went through different
random assortment stages, or through
recombination that mixes different parts of
homologous chromosomes in meiosis.
On the other hand, if the mom’s gametes do go through meiosis, then the haploid egg has to be manipulated so that it is once again diploid. This is called automixis (auto = self) and comes in a couple of flavors. Two eggs can fuse, each being haploid, to produce a diploid super egg. Another way is for the egg to start to develop, go through a few divisions to form what is called a blastomere, and then two blastomere cells will fuse. Finally, a haploid egg or blastomere cell can fuse with a polar body, one of the meiotic products that was cheated of some cytoplasmic factors and did not become a full fledged egg.

The result of any of these fusions is the same, a diploid egg that can develop into a whole organism without fertilization. BUT---- they are not equal to the apomictic egg described above. In meiosis, there is a division of chromosomes and possible recombination to form new sequences. Therefore, no two eggs will have exactly the same DNA, even if produced at the same time by the same mom.

If you fuse these two different eggs (or blastomeres), the sets of chromosomes ARE NOT the same, so even though the offspring will have only maternal DNA, they will not be exact clone of the mom. Automictic parthenogens are therefore called half clones; apomictic parthenogens are full clones.

The fly in the ointment here is sex determination. It is possible for a clonal offspring of a parthenogenic mom to be of the opposite sex – weird enough for you? It all depends on the system that the particular group of animals uses to determine sex.

In mammals, the sex determination system is XX/XY. Females don’t have a Y, so even if by some miracle a mammal could give birth parthenogenically (it doesn’t happen, see below for why), the offspring could be only female. In other animals, this is not so.

There are different sex determination systems in different
groups of animals. The difference between human and
insects is that in humans males have two different sex
chromosomes, while in insects, the male just gets one copy
of the only type of sex chromosome. In the komodo dragon,
the sex determination system is the same as in birds – that
makes sense, birds and reptiles have a common ancestry.
Going back to last week’s example of the Komodo Dragon, their sex determination system is ZZ/WZ, with ZZ = male. Therefore, the automictic fusion of a Z egg with a W egg could produce a female, while two Z eggs fusing would produce a male. On the other hand, apomictic parthenogenesis could produce only males, a Z egg doubles to become a ZZ egg, but a WW egg is not viable.

So komodos would produce more males than females, and their wild populations bear this out. Communities of Komodos can be up to 75% male. It would seem that this is an evolutionary strategy to help the Komodos colonize new islands. Say a female carjacks a log and lands on a new island. She undergoes parthenogenesis because no males are around, and produces males and a few females. Since parthenogenesis is quick (no time wasted on mating and seasonal fertile times) they can build a presence on the island quickly.

Then sexual reproduction can take over, increasing the genetic diversity of the species (because some drift and mutations will have taken place in the offspring). Now the Komodos might be more likely to survive an environmental change that would put on pressure for adaptation.

Another sex determination system is the XX/XO system of many insects. Pea aphids use this system, where XX = female, but those with only one X are male. Parthenogenesis in aphids can also produce only females. And wouldn’t you know it, there are terms for each. If only females are produced, it is called thelytoky; if only males, arrhenotoky, and if both can be produced, deuterotoky.

Most hymenopterans (bees, wasps) are haplodiploid.
This means that the two sexes have different number of
chromosomes. All the males are the product of unfertilized
haploid egg development, but despite this they are sexually
mature. The females are the result of sexual mating and
fertilization of haploid eggs to make them diploid. Even so,
only the rare diploid egg gets the right environment to
become a new queen.
I said above that there is an exception to haploid eggs being worthless, and here it is. Bees are haplodiploid in their sex determination. Males develop from haploid, unfertilized eggs, while females develop from fertilized, diploid eggs. The queen will mate with one or more males to produce new eggs that will be female, while she will lay unfertilized eggs to produce males, In some cases, the female workers will produce unfertilized eggs to become males as well. This is an example of arrhenotoky.

O.K., we’re almost through the terminology, one more set still to get through. Some animals, like the komodos and the pit vipers we have talked about, reproduce through sexual means, but can also reproduce by parthenogenesis under special circumstances. This is called facultative parthenogenesis. On the other hand, some species have abandoned sex all together and ONLY reproduce by parthenogenesis. This is called obligate parthenogenesis and their populations consist of only females – can you imagine the amount of gossip that must go on.

The vast majority of species that have completely abandoned sex (obligate parthenogens) are polyploid. Whiptail lizards are a good example. Of the all the species of whiptails, parthenogenic and sexual, 15 species are obligate parthenogens. And of these, all are polyploid.

Polyploid whiptails have trouble segregating chromosomes because of the increased number of them, and their spindle apparatuses are usually screwed up. If meiosis is going to fail, why use it? And if you aren’t going to use meiosis, why mate with males to produce embryos, just do it yourself? In addition, these species do tend to be found in extreme climates, where males finding females would be more difficult. Parthenogenesis is a way to keep the species going.

In genomic imprinting, genes from mother and dad are
differently regulated. Only one will be active in the
embryo, so you need inputs from ma and pa. The
silencing of the genes in one sex often are the result of
adding methyl groups to the cytosine or adenine bases,
so that they cannot be transcribed into mRNA;
therefore no protein is made.
Facultative parthenogens only resort to asexual reproduction under certain circumstances, usually when males are in short supply, or when increasing numbers quickly is in the species best interest. I say this is usually true, but the paper we talked about last time concerning pit vipers showed that they use parthenogenesis even when males are present. May be they are just fed up with men.

What is common to facultative parthenogens is a lack of genomic imprinting, ie. there are not specific genes provided ONLY by the mom and other genes provided ONLY by the dad. If genes of the different parent must interact to work properly, this is one type genomic imprinting. If the genes exist in both sperm and egg, but one or the other is always silenced, this another type of imprinting.

If there is no genomic imprinting, an individual can survive with just the genes from one parent. However, imprinting is an important regulatory mechanism in all mammals, so we won’t be adopting parthenogenesis any time soon.

Mammals are the only group of animals in which we find genomic imprinting. Of course there is an exception- the monotremes, the platypuses and echidnas. But they’re known for being difficult to put into any one box. They’re mammals, but they lay eggs for gosh sakes!  In fact, it’s their egg laying that negates their necessity for imprinting.

A 2013 review paper looks into the evolution and mechanisms of genomic imprinting in mammals. The imprinted genes are largely involved in transfer of nutrition from the mother to the embryo, ie. the placenta. All mammals have a placenta of one type or another, but monotreme placentas are very short lived, just until the yolk sac forms.

The only surviving monotremes are the platypus and
four species of echidnas. They both look like science
projects gone horribly wrong. They both are mammals,
but they lay eggs. The platypus male has poison spikes
on its hind feet, but its bill is not like a bird bill. The
mouth is on the underside. The echidna has spines and
a long, narrow snout that house both nose and mouth.
Both monotremes have electrosensors in their
bills to find prey.
On the other hand, marsupial mammals (kangaroos, etc.) give birth to very immature young, which then grow bigger and stronger in their pouches. But while in utero, they are still tethered to mama by a placental connection. This makes sense, since monotremes diverged from placental mammals long before the marsupials did.

Since the placenta is so short-lived in monotremes, many of the reasons for imprinting of genes (placental nutrition) are not required. This would leave them free to pursue parthenogenesis as a reproductive strategy, but I am not aware of any documented instances of this.

Genomic imprinting is much more involved than we have described here, and it is involved in more processes than just placental function, including the size of offspring and the competition between males for female eggs. It’s the reason that ligers are so much bigger than tigons! I encourage you to read more about it.

Next week we can look at some very interesting examples of facultative and obligate parthenogenesis, and then some exceptions as to how parthenogenesis works. Exceptions to an exception!

Renfree, M., Suzuki, S., & Kaneko-Ishino, T. (2012). The origin and evolution of genomic imprinting and viviparity in mammals Philosophical Transactions of the Royal Society B: Biological Sciences, 368 (1609), 20120151-20120151 DOI: 10.1098/rstb.2012.0151

For more information or classroom activities, see:

Genomic imprinting –


  1. Hi!
    There's something that's not entirely clear from this post (which I find quite interesting anyway, so thanks!), as you wrote:
    "... On the other hand, apomictic parthenogenesis could produce only males, a Z egg doubles to become a ZZ egg, but a WW egg is not viable..."

    But if apomictic offsprings are FULL clones of their mommy (produced by mitosis) then how is this possible? i.e. all of them should be only females, shouldn't they?


    1. It would seem that way, but we are biased by thinking in terms of XX/XY sex determination, like in humans. But in many species, it is the females that have different sex chromosomes, so when a Z gamete duplicates itself, it can only be ZZ (male), and when a W gamete duplicates, it is non viable, so only a male Komodo can be born parthenogenically by apomixis.

      Remember that apomixis means no mixing of alleles by recombination or mixing of two gametes that have undergone different random assortments in meiosis. That is automixis, and could produce females, if one of the merged gametes was W and one was Z. But in that case, the mixing means that you lose the full clone - it is only a half clone - the sets of chromosomes in two gametes WILL NOT be the same.

  2. Thanks for the answer, the automictic part was clear for me. I think this APOmixis confuses me a bit. I've thought that during APOmixis a diploid egg is produced by the female (ZW) comodo via mitosis - if that's it, there's no meiosis, no recombination and the egg will be the exact clone (also ZW) of its mother, thus being a female.
    Or am I missing something...?


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