Showing posts with label egg. Show all posts
Showing posts with label egg. Show all posts

Wednesday, January 14, 2015

Everybody In The Gene Pool - Plants That Swim

Biology concepts – botany, taxonomy, alternation of generations, cycad, gametophyte, sporophyte, gametes, motility, ginkgo, archegonium, antheridium


In the LOTR The Two Towers we find the tree herders
that can move on their own despite being plants. Today’s
exception is a version of this, if on a much smaller scale.
There are a few types of trees that have motile parts; they
don’t rely on wind, gravity, insects or anything else to
move from one place to another.
Plants are divided up into many categories, and few people agree completely on the groupings. I’ve got a new grouping – swimming plants versus non-swimming plants! It’s as good as anyone else has come up with – let’s investigate the plants that can move on their own.

Plants are most often called by their binomial system names (genus, species), but this is just naming, not categorizing in a large way. Animals, protists, fungi, etc. are all divided into different phyla based on their similarities ad differences. But for some reason (I hope there’s a reason) botany uses divisions instead of phyla.

There are 10-12 divisions of plants (we’ll use 10), covering everything from the mosses to the flowers. Compare that to 21 phyla of animals and it seems like the plants will be easier to classify – but not so fast.

Group plants according to various characteristics and you start to muddy the waters. You might divide them up according to whether they have vascular tissue or not. Non-vascular plants are short because they don’t have vessels to move water very high – these are the Anthocerotophyta (hornworts), the Bryophyta (mosses), and the Marchantiophyta (liverworts). Together they are called the Bryophytes.

The vascular plants are all seven of the other divisions – the Lycopodiophyta (the spikemosses and clubmosses), Pteridophyta (ferns and horsetails), Coniferophyta (conifers), Cycadophyta (cycads), Ginkgophyta (just one species, Ginkgo biloba), the Gnetophyta (a weird group), and Angiospermae (flowering plants).  The conifers, cycads, ginkgo and gnetophytes are often grouped together as the gymnosperms – you’ve probably heard of them.

On the other hand, you might divide the plants up into the seed plants and non-seed plants. In that case, you lump the club mosses and the ferns with the bryophytes, since they all reproduce using spores, not seeds.


Every plant alternates between versions of itself that are
diploid (sporophyte) and haploid (gametophyte). Different
types of plants spend different amounts of time as one or
the other. Mosses and other bryophytes are almost always
in their gametophyte state, while trees are sporophytes with
microgametophytes (antheridium and archegonium) located
in their flowers only.
Lest you think that all those divisions that are seed plants are the same, you can divide them all differently based on whether their seeds are naked (the gymnosperms) or those with fruit (parts of the ovary or accessory organs that overgrow – all in the angiosperms).  Or you could divide them on the basis of how many leaves the embryonic plants have; monocots have one while dicots have two. Angiosperms play by these rules, but gymnosperm seeds don’t, their embryos may have none, one, two, or dozens of cotyledons (embryonic leaves).

Here’s one classification method you may not have heard before – gametophyte dominant vs. sporophyte dominant plants. This has to do with the cycle of life of plants.

Every plant has two lives. Part of its life is spent as a haploid gametophyte (produces haploid gametes) while another plant of the species is a diploid sporophyte.  The sporophyte produces spores that grow into the gametophyte, then the gametophyte produces gametes that join together during fertilization to become a new sporophyte.

Some plant types (like bryophytes) exist mostly in the gametophyte stage and are therefore called gametophyte dominant. Other plants (like trees and flowers) spend all there time as sporophytes and only small parts of them become gametophytes (like pollen or cones).

One way you shouldn’t classify plants is based on their movement. Sure, some plants can grow in a certain direction, toward or away from some stimulus (tropisms, see this post), but plants aren’t motile. They don’t pick up and move themselves from one place to another under their own power.


The Himalayan Balsam is an invasive viny flower that has
become a problem in Europe and is invading the US as well.
Their seed pods contain fins under great strain when fill
with water. The slightest provocation will cause them to
explode, sending seeds more than 25 ft (7.5 m).
Plants also have many ways of moving their seeds and this is sort of a way of plants moving, but I think it’s cheating a bit. Seed dispersal mechanisms can rely on the wind (maples, dandelions) or water (cranberries, coconuts). They can use animals that eat them (many), or just grab ride on them (devil’s claws), or they can burst out (peas) or be shaken out (poppies) and use gravity. But this isn’t really a plant moving by it’s own power.

Plants disperse seeds as new plants, but they also disperse their male gametes in order to find the egg on another plant of their species. You have to get the pollen of seed plants (containing male gametes) or the male gametes themselves to the egg. Like seeds, pollen grains can be moved by insect, by wind, by rain, etc. These are the ways most plants get their male gametes to the egg in order to create a new plant, either in a seed or without a seed.

But there are exceptions, and this is weird exception. Some plants have male gametes that are motile. They swim to the egg! No big deal for animals, they pretty much all have motile male gametes (we’ll look at the exceptions to that), but it’s quite the stunner in plants.


Who knew SpongeBob cartoons were science lesson. Plankton
is green, a phytoplankton – an algae to be precise. Those two
antennae? Probably his two flagella, the way he would swim
around. The feet and the single eye-not so scientific.
Algae are probably the ancestors of all land plants, and we know their gametes have flagella for swimming from our post on them. But some land plants retain this method of male gamete dispersal, but they do include some weird twists. The plants that have motile male gametes cross many of the classes that we described above. There are some seed plants and some spore plants, some vascular and some non-vascular, some are gametophyte dominant and a few are sporophyte dominant – but no flowering plants do this.

Bryophyte males gametes are swimmers. The mosses, liverworts and hornworts  live close to the ground and must have standing water for the make gamete to reach the female gametophyte and egg. The haploid gametophytes are the moss that we usually see. The antheridium grow on the top of the male plants to produce male gametes, while the archegonium on the female plant tip produces the ovule with the egg. When the water is high enough the antheridium releases the male gametes and they swim to the egg using two flagella. The sporophyte (diploid) plant grows from the top of the female gametophyte.

Ferns, horsetails, and club mosses are taller than mosses because they are vascular, but they still require water for their male gametes to swim to the egg. The gametpophyte is a heart shaped leaf that lies near the ground. At one spot the archegonium grows the egg, while the male gametes in the antheridium grow nearby. Water resting on the leaf allows the male gametes to swim across the leaf to the egg (or from leaf to leaf). The sporophyte grows from the heart-shaped gametophyte and is the fern we usually think of.


Cycads (left) are gymnosperms whose trunk are formed from
the bases of the leaves as they grow and are lost. There are
about 300 species of cycads known, with several added every
year. The Ginkgo biloba is the only extant species of the
division. Because its wood is insect resistant, some trees may
be over 2500 years old.
Angiosperms and other seed plants use pollen to send the male gametes to the ovule. When the pollen reaches the archegonium, a tube grows into the ovule and to the egg. The male gamete cells are carried along by the pollen tube right to the front door of the egg. This is when fertilization occurs.  But not all seed plants work this way. A few of the gymnosperms still use a modified version of swimming to the egg.

Cycads (about 300 species) and the lone extant ginkgo, Ginkgo biloba, do have pollen grains that represent the male gametophyte plant. They get blow or carried to the female gametophyte cone and then it gets weird.

The ovule produces a drop of liquid that sticks into the air. The pollen gets caught in this drop and then the drop and the pollen are pulled back into the ovule. The pollen tube grows into the female reproductive organ, but not right to the egg. When the pollen tube reaches the entrance of the archegonium, it ruptures and the gametes are released into a watery fluid that surrounds the eggs.


Cycad and ginkgo male gametes move on their own, the only
exceptions in the seed plants. They have hundreds to thousands
of cilia, as opposed to flagella in bryophyte gametes, which
pull the cell forward. Ginkgo male gametes are huge (0.3 mm),
larger than an entire Wolffia globosa plant.
The male gametes have about a thousand of cilia (not flagella) that pull the cell through the watery environment inside the ovule toward the egg. Fusion and fertilization occurs when the male gametes find the egg – as always. The cycad and ginkgo male gametes swim, but they swim in the indoor pool, not out in the old swimmin’ hole like the bryophytes.

The male gametes of cycads and ferns are very different, from where they swim, to their relative sizes – ginkgo male gametes are HUGE compared to those of ferns, to the use of thousands of cilia as opposed to a couple of flagella.  However, research shows that they are remarkable similar in structure and function.

A 2006 study looked at the proteins involved in gamete movement in ferns and ginkgo. Their results indicated that most of the proteins in both were homologous enough that it suggested a direct descent from bryophyte to gymnosperm, not a case of parallel evolution.


In Guam, there is a neurologic disease that looks a lot like
Alzheimer’s. Research in 2004 found that it was actually
coming from cycad trees. Here’s how it happens. Cyanobacteria
live in the tree roots and put the toxin BMAA into the tree
tissues. Bats eat the fruits and people eat the bats. Than BMAA in
the brain causes disease.
One more cycad exception while we’re here. The cycads can do something that I thought was reserved only for philodendrons and a few relatives. These thermogenic plants can raise the temperature of their male cones by several degrees when the pollen is mature. A 2013 study showed that they can raise the temperature of male cones 2-15 degrees above ambient temperature.

This is believed to attract more insects as pollen distributors, and the researchers did find that more insects visited the plant when the temperature was increased. The mechanism may involve volatilizing more attracting chemicals though the added heat, which would then attract more pollinators (usually weevils).  Pretty advanced for a plant with a so-called primitive reproduction mechanism.

Next week – the base of the undulipodia has a special story all its own. Is it another instance of bacteria evolving into one of our organelles? And it has two very different jobs – which came first?



Suinyuy, T., Donaldson, J., & Johnson, S. (2013). Patterns of odour emission, thermogenesis and pollinator activity in cones of an African cycad: what mechanisms apply? Annals of Botany, 112 (5), 891-902 DOI: 10.1093/aob/mct159

Vaughn, K., & Renzaglia, K. (2006). Structural and immunocytochemical characterization of the Ginkgo biloba L. sperm motility apparatus Protoplasma, 227 (2-4), 165-173 DOI: 10.1007/s00709-005-0141-3

Murch, S., Cox, P., & Banack, S. (2004). A mechanism for slow release of biomagnified cyanobacterial neurotoxins and neurodegenerative disease in Guam Proceedings of the National Academy of Sciences, 101 (33), 12228-12231 DOI: 10.1073/pnas.0404926101


For more information or classroom activities, see:

Alternation of generations –

Seed dispersal –

Pollen –

Cycads –

Ginkgo biloba –

Wednesday, February 27, 2013

The Yolk’s On You

Biology concepts – parthenogenesis, avian reproductive system

Some people practice a form of vegetariansim called veganism. The definition of vegan can be different from person to person, but generally it means that one does not consume or use animal products in which an animal was harmed to obtain them.

Vegans range from those who won’t eat animals or
products that require animals to be harmed, to those
that will not use any animal product whatsoever.
Plants make up their entire diet. How shocked will
they be to learn that research has uncovered that
plants might well have feelings and sensations.
But there are gray areas. Take chicken eggs for instance. Most chicken eggs laid by working hens all over the world are not fertilized. We will talk much more about this in a couple of paragraphs.

Unfertilized eggs can’t develop into a chick. Because of this, some vegans will eat chicken and quail eggs that are certified unfertilized. Now for a potential problem – chickens, quails, turkeys, and other birds are known to undergo parthenogenesis! Unfertilized eggs might have a partially developed embryo inside them. What is a vegan to do?!

The thought that birds might be able to undergo parthenogenesis is not that strange. Reptiles are the ancestors of bird species, and reptiles are famous for their number of parthenogenic species, both obligate and facultative.

The difference between parthenogenesis in birds and in reptiles is that the bird form rarely, very rarely, ends with a chick hatching from the egg. Most die at some point in development - usually early.

The first species in which bird parthenogenesis was studied was the Beltsville Small White (BSW) Turkey. It was recognized in the 1950’s that this breed had some eggs that had started to develop even though they had not been mated to a male.

The researchers then embarked on a long breeding project in which they increased the number of parthenogenic embryos. By breeding females that had a higher tendency to lay parthenogenic eggs  to males from mothers who were more likely to develop parthenogenic eggs, the scientists developed the breed for parthenogens that developed longer and longer.

In some cases (still less than 1%) the parthenogenic eggs would hatch. And some of those turkeys matured fully and lived to a ripe old age! The breed still exists and is still studied, so parthenogenic tom turkeys are born to these females every once in a while.

Zebra finches are native to central Australia, but have
been introduced into North America, Brazil, and
Portugal. They are good for research because they can
breed all year round, usually after strong rains, they
have clutches of five-seven eggs, and they are fast maturing.
Remember that they must be toms because of the sex determination system of birds. Males are ZZ, while their mothers have ZW sex chromosomes. By producing a diploid ovum (egg cell), the moms would double their own DNA, giving either a ZZ or a WW, but WW’s are nonviable. Therefore, all parthenogens would have to be male.

After studies in turkeys were publicized, it was recognized that parthenogenic development was also occurring in chickens. There was a single report of parthenogenesis in a pigeon. Then in the late 2000’s, parthenogenesis in both Chinese Painted Quails and Zebra Finches was recognized. It is important to note that in these later named species, only one parthenogen was noted to survive; that being a chicken in the 1970’s. At that time, molecular methods of genome identification were not available, so we are not sure if this was a true parthenogen.

The other point to note is that these are all domesticated, captive birds. We don’t know if parthenogenesis takes place in birds in the wild. Similar to the cases in other animals, we first recognized parthenogenesis in sharks and komodo dragons in zoos, because that is where people could control if females were exposed to males. Many assumed that parthenogenesis was caused by a lack of males and that they would not give birth from unfertilized eggs in the wild. We now know that isn’t true for komodos, and we have the report showing that pit vipers will undergo parthenogenesis in the wild, even if males are present. Who knows if this is the case for birds.

To understand parthenogenesis in birds, it would help to look at how eggs are produced; we’ll use the chicken as a model. Some weird things can happen with chicken eggs and their process of production is responsible for most of these oddities.

This isn’t the most pleasant of pictures, but it shows the
reproductive tract well. The yellow orbs on the left are the
ova with developing yolk in the ovary. The bigger ones will
be released first. The magnum is larger and adds albumen;
the isthmus is narrower and adds the membranes. Guess
what the shell gland does. The finished eggs exit via the vent.
First off, think of the laying of an egg as the equivalent of a human female’s menstrual cycle. Each month, a woman of child-bearing years will release a mature egg from her ovary (maybe two). In humans, the ovaries trade off each month, but in chickens, only the left ovary and oviduct are functional; the right is there when born, but degenerates over time.

If the human egg is fertilized, the embryo will implant into the wall of the uterus and the placenta will develop. If not, the uterine environment will flush itself out each month and the cycle will begin again. This is different in chickens. Whether the egg is fertilized or not, the ovum (and attached yolk) will be sent on along the oviduct and an egg will be formed.

Chicks are born with 13,000-14,000 ova and they produce no more. Not all will be laid as eggs, but every 26 hours or so, a new ovum with developed yolk (fatty nutrients for the developing chick) will be released from the ovary. The timing of the release is actually controlled by the laying of the egg. When an egg is laid, a new ovum will be released about 30-60 minutes later.

Ovulation is also controlled by the amount of sunlight in the day. Summer day lengths stimulate ovulation, so egg producers manipulate the lights so the hens always think it is summer.

Of course, there is an exception to this. Chickens won’t ovulate after about 3:00 pm! They must where watches. And the entire process for laying an egg takes about 26 hours. This is longer than a day – duh - so each day the chicken will ovulate about 2 hours later. This keeps up until she would be due to ovulate after 3:00. In this case, she just won’t do it, and will wait until the next morning to ovulate. As a result, a chicken will not lay an egg once every six days or so.

Double yolks aren’t really that uncommon, occurring
in about 1in 1000 eggs. However, they are usually
caught in the production process and used for other
egg products instead of putting them in your Styrofoam
box. Quadruple yolks are much less common, but they
do occur, and some breeds are more likely to give
them than others.
When ovum and mature yolk exit the ovary, they enter the oviduct. The first portion is called the infundibulum, and is where the ovum would be fertilized, if a rooster has been in the hen house recently. Whether or not it has been fertilized, the egg then passes into the magnum, which is about 4 inches or so long. In the magnum, the albumen is added around the yolk and ovum. The albumen is clear and provides protection and nutrients to the embryo. It is 90% water and about 10% protein.

After the magnum is the isthmus. This is where the egg is surrounded by the inner and outer membranes. The next stop is the shell gland, and you can guess what is added here. The calcium carbonate shell takes about 20 hours to form around the egg, so this is where the egg spends the majority of its time. Then it is laid by being squeezed out with muscle movement.

Like I mentioned above, weird things can happen during this 25-27 inch trek through the chicken. Sometimes two ova may be released at once. These can both be surrounded by a single albumen and shell and come out as a double yolk egg. There are instances where one or both yolks may be fertilized, but the lack of space and nutrients usually leads to at least one of the chicks dying in the shell, and usually both. The record is nine yolks in a single egg!

On a different note, when the hen is young no ovum may be released, and a small piece of loose tissue could be mistaken for an ovum. In this case, it will be wrapped in albumen, membranes and shell, and a yolkless egg will be produced. I have a student who is seriously considering investigating a way to manipulate chickens to give yolkless eggs all the time – could be a million dollar idea.

There was a recent story that illustrated one more weird possibility. If the muscular movement shoots the egg backward instead of out, it can happen that the developed egg will go through another round of the process. It can also meet up with the next developing ovum. In this case, the developed egg could be surrounded with more albumen, membrane, and then be wrapped in another shell - an egg within an egg! Don’t believe me? Watch the video.

In parthenogenesis, the ovum + yolk will be diploid, the result of endomitosis or fusion of two ova. They will be sent along the path of egg production, and once laid, they look like regular eggs. The embryo will not develop beyond three days or so, so they are hard to tell from unfertilized eggs or those eggs that are fertilized and undergo spontaneous early embryonic death. You probably wouldn’t know if you were eating one.

The Beltsville Small White turkey was developed in the
1930’s and quickly became the most popular turkey
on American plates. The name comes from the USDA
research farm n Beltsville Maryland, where they were
developed. However, in the 1940’s the broad breast
turkey was bred and the BSW quickly faded, except for
in research – they have high rates of parthenogenesis!
That should keep them popular.
The stimulus for diploid egg production is not known; however, the increase in parthenogenesis in the BSW turkeys after breeding them indicates that there is a genetic component to the development of unfertilized eggs. What that component might be is up for grabs. Maybe the breeding selected for females that have an odd hormone profile, or are more apt to undergo endomitosis in their gametes, or ….. you find out and get rich.

In the BSW turkeys, breeding led to later development and finally some live hatchings. This is now being tried in quails as well. Dr. C.D. McDaniel at Mississippi State University is investigating the idea that parthenogenic development actually reduces the hen’s ability to hatch fertilized eggs.

After nine generations of cross breeding females and males to increase parthenogenic development, McDaniel reported in late 2012 that quail that have more parthenogenic events do indeed have fewer fertilized eggs that hatch and develop to mature quails. Late embryonic death decreased, but early death increased dramatically. This is a significant economic question, as it would seem that lower rates of parthenogenesis will lead to greater production of quails.

Next week, we will see that parthenogenesis is not always the “choice” of the female. Sometimes, parthenogenesis can be forced on an animal.

Parker, H., Kiess, A., Robertson, M., Wells, J., & McDaniel, C. (2012). The relationship of parthenogenesis in virgin Chinese Painted quail (Coturnix chinensis) hens with embryonic mortality and hatchability following mating1 Poultry Science, 91 (6), 1425-1531 DOI: 10.3382/ps.2011-01692