Seed Oddities: Vivipary

Seeds house and protect infant plants. When released from their parent plant, they commence a journey that, if successful, will bring them to a suitable location where they can take up residence (upon germination) and carry out a life similar to that of their parents. Their seed coats (and often – in the case of angiosperms – the fruits they were born in) help direct them and protect them in this journey. Physical and chemical factors inhibit them from germinating prematurely – a phenomenon known as dormancy. Agents of dispersal and mechanisms of dormancy allow seeds to travel through time and space — promises of new plants yet to be realized.

There is rarely a need for a seed to germinate immediately upon reaching maturity. In many cases, such as in temperate climates or in times of drought or low light, germinating too soon could be detrimental. The most vulnerable time in a plant’s life comes when it is a young seedling. Thus, finding the right time and space to get a good start is imperative.

The fruits (and accompanying seeds) of doubleclaw (Proboscidea parviflora) are well equipped for long distance dispersal. (via wikimedia commons)

In rare instances, dispersal via seeds offers little advantage; instead, dispersal of live seedlings or propagules is preferable. For this select group of plants, vivipary is part of the reproductive strategy. In vivipary, seeds lack dormancy. Rather than waiting to be dispersed before germinating, viviparous seeds germinate inside of fruits that are still attached to their parent plants.

Occasionally, seeds are observed germinating inside tomatoes, citrus, squash, and other fruits; however, these fruits are usually overripe and often detached from the plant. In these instances, what is referred to as “vivipary” is not a genetic predisposition or part of the reproductive strategy. It’s just happenstance – a fun anomaly. The type of vivipary discussed in this post is actually quite rare, occurring in only a handful of species and prevalent in a select number of environments.

There are three main types of vivipary: true vivipary, cryptovivipary, and pseudovivipary. In true vivipary, a seed germinates inside the fruit and pushes through the fruit wall before the fruit is released. In cryptovivipary, a seed germinates inside the fruit but remains inside until after the fruit drops or splits open. Pseudovivipary is the production of bulbils or plantlets in the flower head. It does not involve seeds and is, instead, a form of asexual reproduction that will be discussed in a future post.

True vivipary is commonly seen among plant communities located in shallow, marine habitats in tropical or subtropical regions, such as mangroves or seagrasses. The term mangrove is used generally to describe a community of plants found in coastal areas growing in saline or brackish water. It also refers more specifically to the small trees and shrubs found in such environments. While not all mangrove species are viviparous, many of them are.

Seedlings of viviparous mangrove species emerge from the fruit and drop from the plant into the salty water below. From there they have the potential to float long or short distances before taking root. They may land in the soil upright, but often, as the tide recedes, they find themselves lying horizontally on the soil. Luckily, they have the remarkable ability to take root and quickly stand themselves up. Doing this allows young plants to keep their “heads” above water as the tides return. It also helps protect the shoot tips from herbivory.

Viviparous seedlings emerging from the fruits of red mangrove (Rhizophora mangle) via wikimedia commons

Another example of vivipary is found in the epiphytic cactus (and close relative of tan hua), Epiphyllum phyllanthus. Commonly known as climbing cactus, this species was studied by researchers in Brazil who harvested fruits at various stages to observe the development of the viviparous seedlings. They then planted the seedlings on three different substrates to evaluate their survival and establishment.

Epiphyllum phyllanthus is cryptoviviparous, so the germinated seeds don’t leave the fruit until after it splits open. In a sense, the mother plant is caring for her offspring before sending them out into the world. The researchers see this as “a form of parental care with subsequent conspecific [belonging to the same species] nursing.” Since the plant is epiphytic – meaning that it grows on the surface of another plant rather than in the soil – local dispersal is important, since there is no guarantee that seeds or propagules dispersed away from the host plant will find another suitable site. That being said, the researchers believe that “vivipary involves adaptation to local dispersal,” since “the greater the dispersal distance is, the higher the risk and the lower the probability of optimal dispersion.”

Epiphyllum phyllanthus via Useful Tropical Plants

While some viviparous seedlings of mangroves can travel long distances from their parent plant and don’t always root into the ground immediately, they maintain their advantage over seeds because they can root in quickly upon reaching a suitable site and lift themselves up above rising tide waters. As the authors of the Epiphyllum study put it, vivipary is “a reproductive advantage that, in addition to allowing propagules to root and grow almost immediately, favors quick establishment whenever seedlings land on suitable substrates.”

There is still much to learn about this unusual and rare botanical feature. The research that does exist is relatively scant, so it will be interesting to see what more we can discover. For now, check out the following resources:

Also, check out this You Tube video of :

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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!

Book Review: What Weeds Are Thinking

Plant intelligence is a burgeoning field of research. Despite being predominantly sessile organisms, plants are able to sense their surroundings and make decisions based on environmental cues. In a certain sense, they can see, hear, smell, and remember even though they don’t have eyes, ears, noses, or brains. Not surprisingly, these fascinating findings have spawned books, podcasts, documentariesarticles, etc. The idea that plants could be intelligent beings like us is something that captures our attention and imagination.

For example, if plants are so smart, does this mean that they actually have thoughts? And if they have thoughts, what could they possibly be thinking? In her new book, What Weeds Are ThinkingErica Crockett takes a stab at what a particularly despised group of plants might think if, indeed, they could have thoughts. Weeds are, in Crockett’s words, “the deviants of the plant world.” This book was her chance to imagine what might be going on inside the minds of these deviants, despite the fact that they don’t have minds.Crockett conjures up the thoughts of 21 different weeds. Each thought is accompanied by an illustration by Sarah Ragan Olson. The drawings are charming, but the thoughts that juxtapose them aren’t always so sweet. Perhaps you imagine weeds to be potty mouthed? Well, so does Crockett. That being said, this book is not for kids, nor is it for anyone sensitive to adult words and themes. Each of the weeds in this book varies in its degree of irreverence – not all of them are so crass and some of them are actually pretty mild-mannered – but that’s just what you’d expect from such a diverse group of plants.

Comfrey is offended by cow manure being used as fertilizer and would rather be fertilized by “the decaying corpses of [its] relations.” Ground ivy is embarrassed and offended by inadvertently seeing the ankles of human passersby.  Plantain is apparently into being stepped on, and prostrate spurge is trying to “rebrand” to make itself more appealing and set itself apart from purslane. Cheatgrass, no surprise here, comes across as a big jerk. Originally from Eurasia, it is now a freedom-loving American, “choking out the rights of the natives.”

Most in line with what I would expect a weed to be thinking – especially one found growing in an urban area – is prickly lettuce. Upset after watching a fellow member of its species ruthlessly dug up, it laments: “Neither of us decided to seed down in the deep crack of this suburban driveway. We were blown here…It’s our home…Yet we are hunted.”

Milkweed is quite aware of its role as the sole food source of the monarch caterpillar. It sees how much humans appreciate monarchs, and admonishes us for killing off its kind: “Keep it up, and I’ll take every last one of those delicate darlings down with me.”

Botanical inaccuracies aside – and there are several – this was a fun book. The main appeal for me is that it is plant-themed and, more specifically, weeds-themed. If you follow this blog, you’ll know that pretty much anything involving weeds is going to get my attention. Beyond that, any project that puts plants in the spotlight and gives them a voice (even in a fictitious sense) is worth checking out. This book is no exception.

As a bonus, I asked Erica Crockett what other plants (apart from weeds) are thinking. This was her response: “Probably really precious or intellectual things. I imagine tomato plants are fairly self-important and zonal geraniums are divas. Bougainvilleas are likely social climbers and oak trees are dull, but honest.”

More Weeds-themed Book Reviews on Awkward Botany:

Field Trip: UBC Botanical Garden and VanDusen Botanical Garden

Last week, we found ourselves in Vancouver, British Columbia for a work-related conference put on by American Public Gardens Association. In addition to learning heaps about plant collections and (among other things) the record keeping involved in maintaining such collections, we got a chance to visit two Vancouver botanical gardens. Both gardens were pretty big, so covering the entire area in the pace we generally like to go in the time that was allotted was simply not possible. Still, we were smitten by what we were able to see and would happily return given the chance. What follows are a few photos from each of the gardens.

UBC Botanical Garden

UBC Botanical Garden is located at the University of British Columbia. Established in 1916, it is Canada’s oldest university botanical garden. We saw a small fraction of the Asian Garden, which is expansive, and instead spent most of our time in other areas, including the Alpine Garden, the Carolinian Forest Garden, the Food Garden, and one of my favorite spots, the BC Rainforest Garden. The Rainforest Garden is a collection of plants native to British Columbia, which was the original focus of UBC Botanical Garden’s first director, John Davidson.

fall foliage of redvein enkianthus (Enkianthus campanulatus)

Franklin tree in bloom (Franklinia alatamaha) in the Carolinian Forest Garden

alpine troughs

bellflower smartweed (Aconogonon campanulatum)

cutleaf smooth sumac (Rhus glabra ‘Laciniata’) in the BC Rainforest Garden

the fruits of Gaultheria pumila in the E.H. Lohbrunner Alpine Garden

Himalayan blueberry (Vaccinium moupinense) in the E.H. Lohbrunner Alpine Garden

VanDusen Botanical Garden

VanDusen Botanical Garden is a 55 acre garden that opened in 1975 and is located on land that was once a golf course. It features an extensive collection of plants from around the world accompanied by a series of lakes and ponds as well as lots of other interesting features (like a Scottish Shelter, a Korean Pavilion, an Elizabethan Maze, and more). Our time there was far too brief. The whirlwind tour we joined, led by the education director, was a lot of fun, and if the threat of missing our bus wasn’t looming, we would have been happy to stay much longer.

Japanese anemone (Anemone x hybrida ‘Whirlwind’)

fall color on the shore of Heron Lake

knees of bald cypress (Taxodium distichum) in R. Roy Forster Cypress Pond

witch hazel (Hamamelis x intermedia ‘Pallida’)

a grove of giant redwoods (Sequoiadendron giganteum)

We tried the fruit of dead man’s fingers (Decaisnea insignis). It tastes a bit like watermelon.

Japanese stewartia (Stewartia pseudocamellia)

More Awkward Botany Field Trips:

How to Identify Puncture Vine (a.k.a. the Goathead Monster)

This post originally appeared on Idaho Botanical Garden’s blog. With the first annual Boise Goathead Fest fast approaching, the purpose of this post is to help people in the Treasure Valley identify goatheads so that they can collect them for drink tokens to use at the event. I’m reposting it here in hopes that people around the globe who are tormented by goatheads might benefit from it. All photos in this post were taken by Anna Lindquist.

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If you have spent much time on a bicycle in Boise, chances are you have been the victim of a goathead-induced flat tire. You probably even got a good look at the spiky nutlet as you went to remove it from your tire. But where did the culprit come from? No doubt, it came from a plant. But which one?

This is particularly useful to know right now because the first annual Boise Goathead Fest is coming up, and if you manage to fill a garbage bag full of these noxious weeds before the end of July, you will earn yourself a drink token. Fortunately, this plant is fairly easy to identify; however, there are a few look-a-likes, so it is important to familiarize yourself with the plant in question so you can be sure you are pulling the right one.

puncture vine (Tribulus terrestris)

Puncture vine, also known as goathead or Tribulus terrestris, is a warm season annual that is native to the Mediterranean region of southern Europe. It was introduced to North America unintentionally by early European settlers when the plant’s blasted burs snuck their way across the ocean in sheep wool. Since then, puncture vine has spread across the continent prolifically thanks to the hitchhiking prowess of its seeds.

Behold, the infamous Goathead Monster.

Puncture vine has a prostrate habit, meaning that its branches lie flat on the ground, spreading outward from a central location. It grows upward only when it is being shaded or crowded out. Its leaves are divided into several tiny leaflets, and its flowers are small and bright yellow with five petals. It is an otherwise pretty plant were it not for the threatening, jagged fruits that follow the flowers. As these fruits dry, they dislodge from the plant, split into five pieces, and lay in wait to puncture your tire, work their way into the bottom of your shoe or the foot of an animal, or latch onto some errant fur.

puncture vine (Tribulus terrestris)

Depending on the conditions, puncture vine either remains fairly small or spreads as much as six feet wide. Fruits start forming shortly after flowering, and seeds ripen soon after that, so if the plant isn’t removed quickly – nutlets and all – future populations are guaranteed. Luckily the plants are fairly easy to remove. Unless the ground is particularly compact, they pull up easily, and if they break off at the root, they generally don’t sprout back.

Virtually any plant that has a prostrate growth habit and is actively growing in the summer could, at first glance, be mistaken for puncture vine. Closer inspection will help confirm the plant’s true identity. Two plants that might confuse you are purslane and spotted spurge. Both of these species can be found growing in full sun in disturbed or neglected sites in close company with puncture vine.

Purslane has tiny, yellow, five-petaled flowers similar to puncture vine; however, its leaves are glossy and succulent-like and its stems and leaves often have a red to purple hue to them. Purslane seeds are miniscule, and while the plant can be a nuisance in a garden bed, it poses no threat to bicycles or wildlife.

purslane (Portulaca oleracea)

Spotted spurge, also known as prostrate spurge, can be quickly distinguished by the milky sap that oozes from its broken stems. Its leaves are generally reddish purple on the undersides with a purple spot on top. Its flowers are minute and its seeds even smaller. Because its sap contains latex and other chemicals, it can irritate the skin and poison creatures that dare eat it.

spotted spurge (Euphorbia maculata)

Both of these plants are introduced, weedy species, so even if they won’t count towards your drink token, it still doesn’t hurt to pull them. Puncture vine, however, is included on Idaho’s noxious weed list, which means it is particularly problematic. So take this opportunity to pull as many as you can, and hopefully we can put a sizeable dent in the population of a plant that has tormented Boise bicyclists for far too long.

See Also: Plant vs. Bike

Eating Weeds: Burdock

If we agree that weeds can be famous while simultaneously being infamous, a list of famous weeds must include burdock. Its fame largely comes from being an inspiration for the hook-and-loop fastener, Velcro. The idea for this revolutionary product came when Swiss inventor, George de Mestral, was removing burs – the dried inflorescences of burdock – from his dog in the early 1940’s. Most of us have experienced this, pulling out burs from animal hair or our own clothing, but few have felt inspired to develop a new product. Infamy reigns supreme.

But burdock’s fame isn’t tied to Velcro. Its tenacious, sticky burs, which house the seeds, have been attaching themselves to humans and other animals for centuries, frustrating those who have to remove them but finding new places to grow in the process. And what better way to pay tribute to this phenomenon than to dress oneself in hundreds of burs and parade around town calling yourself the Burry Man? Lest you think I’m crazy, just such a thing has been part of an annual celebration for over 300 years in a town outside of Edinburgh, Scotland.

burs of common burdock (Arctium minus)

Of course, burdock is more than its burs. Other, perhaps less celebrated features, are its edible roots and shoots. Its leaves are also edible, but most people find them too bitter to bother. Green Deane suggests wrapping the leaves around food to cook on a campfire. Both the roots and shoots can be eaten raw or cooked, and the fermented roots along with dandelion roots are traditional ingredients in the British beverage, dandelion and burdock. The roots, shoots, and leaves of burdock have also had a wide variety of medicinal uses.

Two species of burdock have become naturalized in North America – Arctium minus and Arctium lappa. Both species are biennials or short-lived perennials. They start out as rosettes of large leaves with woolly undersides. The leaves grow to a foot or more long and wide. At this stage burdock is similar in appearance to rhubarb. Burdock has a large taproot, which can extend down to three feet in the ground. The taproot continues to grow as the rosette expands. When the plant has reached a certain size it begins to put up a branching flower stalk. It is in the rosette stage, before the plant bolts, that the taproot should be harvested.

As the flower stalk grows, the plant takes on a pyramidal shape, with the leaves along the stalk getting increasingly smaller with height. The plant can reach several feet tall, with one source describing them as towering up to ten feet. The stalks should be harvested before the plants start flowering. Multiple flower heads are produced at the ends of the branching stalk. The inflorescences are composed of purple, tubular, disc florets that are encased and encircled in a series of hooked bracts. The flower heads resemble thistle flowers, but the plant is easy to distinguish from thistles due to its large, soft leaves. Speaking of the leaves, one photographer found them alluring enough to compile a series of photos of them.

Common burdock (Arctium minus): the woolly undersides of the leaves and the tops of the taproots

While burdock can be nuisance plant, it is not particularly noxious. In The Book of Field and Roadside, John Eastman writes, “Burdock cannot be labeled a truly invasive weed, for it rarely intrudes into cultivated fields. Tilling usually controls and eradicates burdock populations. Its favored havens are the disturbed soils of roadsides, railroads, fence rows, vacant lots, and around sheds and old buildings.” In Wild Urban Plants of the Northeast, Peter Del Tredici also comments on burdock’s preference for minimally maintained locations including “vacant lots and rubble dump sites; the edges of emergent woodlands; the sunny borders of freshwater wetlands, ponds, and streams; and on unmowed highway banks and median strips with frequent salt applications.”

I harvested my burdock roots along an unmaintained fence line surrounding a series of raised flower beds. I chose a simple recipe for making burdock chips that involved peeling the roots, cutting them into thin slices, dressing them with olive oil and salt, and baking them in the oven. Since the author of this recipe mentioned buying burdock from a store, they were probably using Arctium lappa, or greater burdock, which is commonly cultivated, especially in Asian countries. Both species can be prepared in similar ways.

burdock roots

The burdock chips had a pleasant nutty flavor, but they were also a little stringy and tough to chew. If I were to do it again, I would probably use a recipe like this one that involves parboiling and then frying. Sierra suggested grating the roots and frying them in bacon grease, which would probably do the trick. There are also recipes for pickled burdock roots, which would be fun to try.

Because the plants I harvested were still in their rosette stage and there weren’t any other plants in the area that were bolting, I didn’t try the shoots. But I’ll keep my eye out, and when I find some I may have to write a part two.

Eating Weeds: Pineapple Weed

When I wrote about pineapple weed (Matricaria discoidea) last year during the Summer of Weeds, I knew that it was edible but I didn’t bother trying it. Pineapple weed is one of my favorite native weeds (yes, it happens to be a native of northwestern North America). I enjoy its sweet fragrance, its frilly leaves, its “petal”-less flowers, and its diminutive size. I also appreciate its tough nature. Now that I have tried pineapple weed tea, I have another thing to add to this list of pros.

pineapple weed (Matricaria discoidea)

One thing about pineapple weed that always impresses me is its ability to grow in the most compacted soils. It actually seems to prefer them. It is consistently found in abundance in highly trafficked areas, like driveways, parking lots, and pathways, seemingly unfazed by regular trampling. Referring to pineapple weed in one of his books about wildflowers, botanist John Hutchinson wrote, “the more it is trodden on the better it seems to thrive.” This is not something you can say about too many other plants.

Both the leaves and flowers of pineapple weed are edible. The flowers seem to be the more common of the two to consume, generally in tea form. In his book Wild Edible and Useful Plants of Idaho, Ray Vizgirdas writes, “A delicious tea can be made from the dried flowers of the plant. The leaves are edible, but bitter. The medicinal uses of pineapple weed are identical to that of chamomile (Anthemis). Used as a tea it is a carminative, antispasmodic, and mild sedative.” In Wild Urban Plants of the Northeast, Peter Del Tredici writes, “A tea made from the leaves has been used in traditional medicine for stomachaches and colds.”

I harvested my pineapple weed at the end of a dirt parking lot and in an adjacent driveway/pathway. I noted how the pineapple weed’s presence waned as I reached the edges of the parking lot and pathway where, presumably, the ground was less compact. Maybe it has more to compete with there – other weeds – and so it shows up less, or maybe its roots simply “prefer” compact soils. Perhaps a little of both. Once I got my harvest home, I rinsed it off and left it to dry. Later, I snipped off the flower heads and made a tea.

I probably used more water than I needed to, so it was a bit diluted, but it was still delicious. It smelled and tasted a lot like chamomile. Sierra agreed. With a little honey added, it was especially nice. Sierra agreed again. The flowers of pineapple weed can be used fresh or dried. They can also be mixed with other ingredients to make a more interesting tea, like the recipe found here.

If you are hesitant to take the leap into eating weeds, a tea may be the simplest thing you can try. Pineapple weed tea is a great way to ease yourself into it. Apart from maybe having to harvest it from strange places, it probably isn’t much different from other teas you have tried, and, from my experience, it’s delightful.