Seed Oddities: Apomixis and Polyembryony

Plants have uncanny ways of reproducing themselves that are unparalleled by most other living things. Offshoots of themselves can be made by sending out modified stems above or beneath the ground which develop roots and shoots (new plants) at various points along the way. Various other underground stem and root structures can also give rise to new plants. Small sections of root, stem, or leaf can, under the right conditions, push out new plantlets in a fashion that seems otherworldly. (Picture chopping off a bit of your finger and growing a whole new you from it.)

These are some of the ways in which plants reproduce asexually, and it’s kind of freaky if you think about it. Plants can clone themselves. But one major disadvantage of reproducing this way is that clonal offspring are genetically identical to the parent plant, which truncates any advantage that might be gained by genetic mixing between two separate plants. For one, it means that a plant population composed of all clones is at risk of being wiped out if something in the environment comes along (such as a disease or change in climate) and none of the plants in the population have adapted any sort of resistance to it.

New plants forming along the lateral stems of Ranunculus flammula – via wikimedia commons

That’s where seeds come in. Seeds are produced sexually, when the gametes of one plant fuse with the gametes of another. Genetic recombination occurs, and a genetically unique individual is born, housed within a seed. Unless, of course, that seed is produced asexually. Then the seed is a clone, and we’re back to where we started.

Apomixis is the process by which seeds are produced asexually. In flowering plants, this means that viable seeds are formed even when flowers haven’t been pollinated. In some cases, pollination stimulates apomixis or is required to produce endosperm; but either way, the result is the same: an embryo containing an exact copy of the genes of its single parent plant.

To understand this process, it’s important to familiarize yourself with the basic anatomy of an ovule, the part of a plant where embryos are formed and which ultimately becomes a seed. In gymnosperms, ovules sit inside cones; in angiosperms, they are surrounded by an ovary. The wall of the ovule is called an integument. A small opening at the top of the ovule, known as a micropyle, is where the pollen tube enters. Diploid cells of the nucellus compose the interior of the ovule, and within the nucellus resides the megasporocyte, which is where meiosis occurs and egg cells are produced. In sexual reproduction, a germ cell introduced through the pollen tube fuses with the egg cell to form a zygote and eventually an embryo. In the case of apomixis, the fusion of germ cells isn’t necessary for an embryo to form.

ovule anatomy via wikimedia commons

There are three main types of apomixis: diplospory, apospory, and adventitious embryony. In diplospory, the megasporocyte skips meiosis and produces diploid cells instead of haploid cells (germ cells). These unreduced cells go on to form an embryo inside of the embryo sac, just like an egg cell would if it had been fertilized with a pollen cell. Additional unreduced cells go on to form endosperm, and the ovule then matures into a seed. This type of apomixis is common in dandelions (Taraxacum officinale). As much as bees love visiting dandelion flowers, their pollination services are not required for the production of viable seeds. Yet another reason you are stuck with dandelions in your yard whether you like it or not.

In apospory, an embryo develops inside of an embryo sac that has been formed from cells in the nucellus. Embryo development within the megasporocyte is bypassed; however, pollination is usually necessary for endosperm to form. Plant species in the grass family commonly produce seeds using this type of apomixis.

Adventitous embryony is also known as sporophytic apomixis because an embryo is formed outside of an embryo sac. Cells from either the integument or the nucellus produce an embryo vegetatively. In this case, a sexually produced embryo can form along with several vegetatively produced embryos. Extra embryos die off and a single, surviving embryo is left inside the mature seed. But not always. Two or more embryos occasionally survive, including the sexually produced one. The mature seed then consists of multiple embryos. This phenomenon is called polyembryony and is common in citrus and mangoes. When it comes to plant breeding, polyembryony is incredibly useful because the asexually derived seedlings are exact copies of their parent, which means the desirable traits of a specific cultivar are retained.

Depiction of seed with three viable embryos after germination.

Polyembryony can occur in a number of ways, and not always as a result of apomixis. In some species, additional embryos “bud off” from the sexually produced embryo. This is called cleavage polyembryony and is known to happen frequently in the pine family (Pinaceae), as well as other plant families. Another common form of polyembryony in gymnosperms is simple polyembryony, in which several egg cells in a single ovule are fertilized resulting in the development of multiple embryos. This doesn’t always mean there will be multiple seedlings sprouting from a single seed. Most embryos usually fail to mature, and only one prevails. However, sometimes more than one survives, and if you’re lucky, you’ll find a seed with multiple plant babies pushing out from the seed coat.

Up Next: Vivipary!

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Inside of a Seed: Gymnosperms

“Every tree has to stay where it put down roots as a seedling. However, it can reproduce, and in that brief moment when tree embryos are still packed into seeds, they are free. The moment they fall from the tree, the journey can begin.” — The Hidden Life of Trees by Peter Wohlleben

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Seed plants – also known as spermatophytes – make up the largest group of plants on earth. Seed plants consist of five divisions, and among them the angiosperm division (a.k.a. flowering plants) dominates in its number of species. The four remaining divisions are referred to collectively as gymnosperms. This incudes the cycads (Cycadophyta), Ginkgo biloba (the only living species in the division Ginkgophyta), gnetophytes (Gnetophyta), and the conifers (Coniferophyta). Conifers are by far the largest and most widespread gymnosperm division.

Angiosperms and gymnosperms have different evolutionary histories, resulting in their distinct genetic and morphological differences. That being said, an overly-simplistic way of differentiating the two groups is to say that, while both groups produce seeds, angiosperms produce flowers and fruits while gymnosperms produce pollen cones and seed cones. There are always exceptions (Ginkgo biloba, for example, doesn’t produce cones), but for the most part, this is the case.

Pollen cones (top) and seed cones (bottom) of mugo pine (Pinus mugo) via wikimedia commons

Sexual reproduction in gymnosperms follows a familiar pattern. Pollen, which contains the male sex cells, is produced in pollen cones, which are essentially miniature branches with modified leaves called scales that house the male reproductive organs. Mature pollen is shed and carried away by the wind. Lucky pollen grains make their way to the female cones, which are also modified branchlets, but are a bit more complex. Scales sit atop bracts, and on top of the scales are ovules – the female reproductive structures. During fertilization, the bracts open to collect pollen and then close as the seed develops.

When pollen lands on an ovule it forms pollen tubes that help direct the male sex cells to the egg cells inside. The process is similar to pollen tubes extending down the style of a flower. In flowering plants, additional pollen cells combine with cells in the ovule to produce endosperm, a storage tissue that feeds the growing embryo. This doesn’t happen in gymnosperms. Instead, haploid cells within the ovule develop into storage tissue and go on to serve the same role.

The ovule eventually matures into a seed, and the cone opens to release it. The seed sits atop the scale rather than enclosed within a fruit, as it would be in an angiosperm. For this reason gymnosperms are said to have naked seeds. The development of seeds can also be much slower in gymnosperms compared to angiosperms. In some species, seeds don’t reach maturity for as long as two years.

Seed cones and winged seeds of mugo pine (Pinus mugo) via wikimedia commons

Seeds in the genus Pinus are excellent representations of typical gymnosperm seeds. Their basic components are essentially identical to the seeds of angiosperms. The seed coat is also referred to as an integument. It was once the outer covering of the ovule and has developed into the seed covering. A micropyle is sometimes visible on the seed and is the location where the pollen cells entered the ovule. The storage tissue, as mentioned above, is composed of female haploid cells that matured into storage tissue in the ovule. Like angiosperms, the embryo is composed of the radicle (embryonic root), the hypocotyl (embryonic shoot), and cotyledons (embryonic leaves).

Angiosperms can be divided into monocotyledons and dicotyledons according to the number of cotyledons their embryos have (monocots have one, dicots have two). Gymnosperms are considered multi-cotyledonous because, depending on the species, they can have a few to many cotyledons.

Seedling of Swiss pine (Pinus cembra) showing multiple cotyledons via wikimedia commons

For the sake of this introduction to gymnosperm seeds, I have offered a simple overview of the production of seeds in the conifer division. Sexual reproduction and seed formation in the other three gymnosperm divisions is a similar story but varies according to species. Even within the conifers there are differences. For example, the “seed cones” of several gymnosperm species can actually be quite fruit-like, which serves to attract animals to aid in seed dispersal. Also, the pollen of gymnosperms is often thought of as being wind dispersed (and occasionally water dispersed in the case of Ginkgo biloba and some cycads); however, researchers are continuing to discover the pivotal role that insects play in the transfer of pollen for many cycad species, just as they do for so many species of angiosperms.

All of this to say that Botany 101 is simply a window into what is undoubtedly an incredibly diverse and endlessly fascinating group of organisms, and that, as with all branches of science, there is still so much to discover.

Inside of a Seed: Two Monocots

“Seeds are travelers in space and time – small packages of DNA, protein, and starch that can move over long distances and remain viable for hundreds of years. These packages have everything they need not only to survive, but also to grow into a plant when they encounter the right conditions.”      The Book of Seeds by Paul Smith

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As illustrated in last week’s post, the mature seeds of dicots – depending on the species – can be either with or without endosperm (a starchy food packet that feeds a growing seedling upon germination). Seeds without endosperm store these essential sugars in their cotyledons. Monocotyledons (or monocots, for short) are a group of flowering plants (i.e. angiosperms) whose seedlings are composed of a single cotyledon. With the exception of orchids, the seeds of monocots always contain endosperm.

The first of two examples of monocot seeds is the common onion (Allium cepa). The embryo in this seed sits curled up, surrounded by endosperm inside of a durable seed coat.

If you have ever sown onion seeds, you have watched as the single, grass-like cotyledon emerges from the soil. The seed coat often remains attached to the tip of the cotyledon like a little helmet as it stretches out towards the sky. Soon the first true leaf appears, pushing out from the base of the cotyledon. The source of this first leaf is the plumule hidden within the cotyledon.

The fruit of plants in the grass family – including cereal grains like wheat, oats, barley, rice, and corn – is called a caryopsis. In this type of fruit, the fruit wall (or pericarp) is fused to the seed coat, making the fruit indistinguishable from the seed. The embryos in these seeds are highly developed, with a few more discernible parts. A simplified diagram of a corn seed (Zea mays) is shown below. Each kernel of corn on a cob is a caryopsis. These relatively large seeds are great for demonstrating the anatomy of seeds in the grass family.

In these seeds there is an additional layer of endosperm called aleurone, which is rich in protein and composed of living cells. The cells of the adjacent endosperm are not alive and are composed of starch. The embryo consists of several parts, including the cotyledon (which, in the grass family, is also called a scuttelum), coleoptile, plumule, radicle, and coleorhiza. The coleoptile is a sheath that protects the emerging shoot as it pushes up through the soil. The plumule is the growing point for the first shoots and leaves, and the radicle is the beginning of the root system. The emerging root is protected by a root cap called a calyptra and a sheath called a coleorhiza.

Germination begins with the coleorhiza pushing through the pericarp. It is quickly followed by the radicle growing through the coleorhiza. As the embryo emerges, a signal is sent to the endosperm to start feeding the growing baby corn plant, giving it a head start until it can make its own food via photosynthesis.

corn seeds (Zea mays)

Up Next: We’ll take an inside look at the seeds of gymnosperms.

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