self pollination

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The Hidden Flowers of Viola

Violas keep a secret hidden below their foliage. Sometimes they even bury it shallowly in the soil near their roots. I suppose it’s not a secret really, just something out of sight. There isn’t a reason to show it off, after all. Showy flowers are showy for the sole purpose of attracting pollinators. If pollinators are unnecessary, there is no reason for showy flowers, or to even show your flowers at all. That’s the story behind the cleistogamous flowers of violas. They are a secret only because unless you know to look for them, you would have no idea they were there at all.

Cleistogamy means closed marriage, and it describes a self-pollinating flower whose petals remain sealed shut. The opposite of cleistogamy is chasmogamy (open marriage). Most of the flowers we are familiar with are chasmogamous. They open and expose their sex parts in order to allow for cross-pollination (self-pollination can also occur in such flowers). Violas have chasmogamous flowers too. They are the familiar five-petaled flowers raised up on slender stalks above the green foliage. Cross-pollination occurs in these flowers, and seed-bearing fruits are the result. Perhaps as a way to ensure reproduction, violas also produce cleistogamous flowers, buried below their leaves.

an illustration of the cleistogamous flower of Viola sylvatica opened to reveal its sex parts — via wikimedia commons

Flowers are expensive things to make, especially when the goal is to attract pollinators. Colorful petals, nectar, nutritious pollen, and other features that help advertise to potential pollinators all require significant resources. All this effort is worth it when it results in the ample production of viable seeds, but what if it doesn’t? Having a method for self-pollination ensures that reproduction will proceed in the absence of pollinators or in the event that floral visitors don’t get the job done. A downside, of course, is that a seed produced via self-pollination is essentially a clone of the parent plant. There will be no mixing of genes with other individuals. This isn’t necessarily bad, at least in the short term, but it has its downsides. A good strategy is a mixture of both cross- and self-pollination – a strategy that violas employ.

The cleistogamous flowers of violas generally appear in the summer or fall, after the chasmogamous flowers have done their thing. The fruits they form split open when mature and deposit their seeds directly below the parent plant. Some are also carried away by ants and dispersed to new locations. Seeds produced in these hidden flowers are generally superior and more abundant compared to those produced by their showy counterparts. People who find violas to be a troublesome lawn weed – expanding far and wide to the exclusion of turfgrass – have these hidden flowers to blame.

That being said, there is a defense for violas. In the book The Living Landscape by Rick Darke and Doug Tallamy, Tallamy writes: “Plants such as the common blue violet (Viola sororia), long dismissed by gardeners as a weed, can be reconstituted as desirable components of the herbaceous layer when their ecosystem functionality is re-evaluated. Violets are the sole larval food source for fritillary butterflies. Eliminating violets eliminates fritillaries, but finding ways to incorporate violets in garden design supports fritillaries.”

sweet violet (Viola odorata)

In my search for the cleistogamous flowers of viola, I dug up a sweet violet (Viola odorata). I was too late to catch it in bloom, but the product of its flowers – round, purple, fuzzy fruits – were revealed as I uprooted the plant. Some of the fruits were already opening, exposing shiny, light brown seeds with prominent, white elaiosomes, there to tempt ants into aiding in their dispersal. I may have missed getting to see what John Eastman calls “violet’s most important flowers,” but the product of these flowers was certainly worth the effort.

Fruits formed from the cleistogamous flowers of sweet violet (Viola odorata)

Up close and personal with the fruit of a cleistogamous flower

The seeds (elaiosomes included) produced by the cleistogamous flower of sweet violet (Viola odorata)

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Summer of Weeds: Eating Purslane

If it wasn’t so prolific and persistent, purslane would probably be a welcome guest in our vegetable gardens and edible landscapes. Easily among the most nutritious and versatile of the edible weeds, Portulaca oleracea is an annoyingly abundant annual that has inserted itself into garden beds and croplands in temperate climates across the globe. Thought to have originated in India or somewhere in Eurasia, purslane invaded North America long before Europeans did and has been naturalized across much of the continent for hundreds of years.

common purslane (Portulaca oleracea)

There are over 100 known species in the genus Portulaca, the only genus in the family Portulacaceae (otherwise known as the purslane family). Common purslane is a succulent plant with paddle- or teardrop-shaped leaves that generally grows low to the ground, forming a thick mat. It reaches for the sky when grown in shade or when competing with other plants for space. It produces little, yellow flowers that only open in bright sun and are typically self-pollinated. A small capsule containing dozens of tiny, black seeds quickly follows each flower. Each plant can produce tens of thousands of seeds, which remain viable for around 40 years.

Attempts to remove purslane by cultivation may only aid its survival. Broken pieces of the plant can take root in the soil, and uprooted plants can re-root if they are in contact with soil. Stirring up the ground brings to the surface seeds from purslane’s extensive seed bank. These freshly exposed seeds can then germinate, taking advantage of disturbance and open space. For all these reasons and more, John Eastman writes in The Book of Field and Roadside: “Purslane knows how to live and linger.”

The ever-urban and ever-common purslane.

The seeds of purslane germinate in late spring and throughout the summer when the soil has reached at least 75 – 80° Fahrenheit. It is adapted to high heat and dry soils. In order to conserve water, it switches to CAM photosynthesis when conditions are particularly hot and dry. In this photosynthetic pathway, carbon dioxide is stored as malic acid during the night and then converted back during the day. This means that, when it comes to eating purslane, the flavor changes depending on when the plant is harvested. In The Wild Wisdom of Weeds, Katrina Blair discusses this phenomenon: “In the morning purslane leaves contain as much as ten times more malic acid, making them very sour tasting. If you prefer a milder tasting purslane, harvest your greens in late afternoon and if you want more zing to your recipes, gather the leaves at dawn.”

Speaking of eating purslane, if all the claims are to be trusted, there may not be a more nutritious weed. In A Feast of Weeds, Luigi Ballerina calls it “a health bomb” because “it contains more omega-3 fatty acids than almost any other green, not to mention vitamins A, B, and C and beta carotene.” Blair calls it “one of the most nutritious plants on Earth,” and goes on to sing praises about its richness in dietary fiber, vitamins, minerals, protein, etc. Funnily enough, in describing the health benefits of purslane, Ballerina also quotes ancient sources claiming that “purslane calms sexual excitement.” Apparently it not only “eliminate[s] sensual dreams, but if used too much, it often extinguishes all ardor and even the capacity to procreate.”

With that caveat in mind, I tried it anyway. I had eaten it before, but nothing more than a leaf here and there and once in a green salad. I picked two recipes to try: Walnut Purslane Coleslaw from The Wild Wisdom of Weeds and Potatoes and Purslane from A Feast of Weeds. I’m generally a big fan of coleslaw, but for whatever reason I found this recipe to be a little bland. It was missing something, but I couldn’t put my finger on it. The purslane seemed to add a vague slimy-ness to it, which it will do on account of its mucilaginous nature.

Walnut Purslane Coleslaw

The Potatoes and Purslane recipe involved cooking the purslane. I enjoyed the finished product both hot and cold. The purslane added a sort of lemon-y spinach flavor. Those who tried it with me also liked it. The potato recipe was made with purslane that had been harvested in the morning, which may explain the strong lemon-y flavor. The coleslaw was made with purslane harvested in late afternoon, which may explain its blandness. I will have to try it the other way around for comparison. Purslane recipes abound in books and on the internet; browsing through them, I am intrigued enough to consider trying others. I think I’ll start with pickled purslane, purslane pesto, and perhaps, purslane sauerkraut.

Potatoes and Purslane

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Do you have a favorite purslane recipe? Share it in the comment section below.

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Ethnobotany: White Man’s Foot, part one

“Plantains – Plantago major – seem to have arrived with the very first white settlers and were such a reliable sign of their presence that the Native Americans referred to them as ‘white men’s footsteps.'” – Elizabeth Kolbert (The Sixth Extinction)

“Our people have a name for this round-leafed plant: White Man’s Footstep. Just a low circle of leaves, pressed close to the ground with no stem to speak of, it arrived with the first settlers and followed them everywhere they went. It trotted along paths through the woods, along wagon roads and railroads, like a faithful dog so as to be near them.” – Robin Wall Kimmerer (Braiding Sweetgrass)

photo credits: wikimedia commons

photo credit: wikimedia commons

Plantago major is in the family Plantaginaceae – the plantain family – a family that consists of at least 90 genera, several of which include common species of ornamental plants such as Veronica (speedwells), Digitalis (foxgloves), and Antirrhinum (snapdragons). The genus Plantago consists of around 200 species commonly known as plantains. They are distributed throughout the world in diverse habitats. Most of them are herbaceous perennials with similar growth habits, and many have ethnobotanical uses comparable to P. major.

Originating in Eurasia, P. major now has a cosmopolitan distribution. It has joined humans as they have traveled and migrated from continent to continent and is now considered naturalized throughout most temperate and some tropical regions. In North America, P. major and P. lanceolata are the two most common introduced species in the Plantago genus. P. major has a plethora of common names – common plantain being the one that the USDA prefers. Other names include broadleaf plantain, greater plantain, thickleaf plantain, ribgrass, ribwort, ripplegrass, and waybread. Depending on the source, there are various versions of the name white man’s foot, and along the same line, a common name for P. major in South Africa is cart-track plant.

P. major is a perennial – albeit sometimes annual or biennial – herbaceous plant. Its leaves form a rosette that is usually oriented flat against the ground and reaches up to 30 cm wide. Each leaf is egg-shaped with parallel veins and leaf margins that are sometimes faintly toothed. The inflorescence is a leafless spike up to 20 cm tall (sometimes taller) with several tiny flowers that are a dull yellow-green-brown color. The flowers are wind pollinated, and the plants are highly prone to self-pollination. The fruits are capsules that can contain as many as 30 seeds – an entire plant can produce as many as 14,000 – 15,000 seeds at once. The seeds are small, brown, sticky, and easily transported by wind or by adhering to shoes, clothing, animals, and machinery. They require light to germinate and can remain viable for up to 60 years.

An illustration of three Plantago species found in Selected Weeds of the United States - Agriculture Handbook No. 366 circa 1970

An illustration of three Plantago species found in Selected Weeds of the United States – Agriculture Handbook No. 366 circa 1970

P. major prefers sunny sites but can also thrive in part shade. It adapts to a variety of soil types but performs best in moist, clay-loam soils. It is often found in compacted soils and is very tolerant of trampling. This trait, along with its low-growing leaves that easily evade mower blades, explains why it is so commonly seen in turf grass. It is highly adaptable to a variety of habitats and is particularly common on recently disturbed sites (natural or human caused) and is an abundant urban and agricultural weed.

Even though it is wind pollinated, its flowers are visited by syrphid flies and various bee species which feed on its pollen. Several other insects feed on its foliage, along with a number of mammalian herbivores. Cardinals and other bird species feed on its seeds.

Humans also eat plantain leaves, which contain vitamins A, C, and K. Young, tender leaves can be eaten raw, while older leaves need to be cooked as they become tough and stringy with age. The medicinal properties of  P. major have been known and appreciated at least as far back as the Anglo-Saxons, who likely used a poultice made from the leaves externally to treat wounds, burns, sores, bites, stings, and other irritations. Native Americans, after seeing the plant arrive with European settlers, quickly learned to use the plant as food and medicine. It could be used to stop cuts from bleeding and to treat rattlesnake bites. Apart from external uses, the plant was used internally as a pain killer and to treat ulcers, diarrhea, and other gastrointestinal issues.

P. major has been shown to have antibacterial, anti-inflammatory, antioxidant, and other biological properties; several chemical compounds have been isolated from the plant and deemed responsible for these properties. For this reason, P. major and other species of Plantago have been used to treat a number of ailments. The claims are so numerous and diverse that it is worth exploring if you are interested. You can start by visiting the following sites:

"White man's footstep, generous and healing, grows with its leaves so close to the ground that each step is a greeting to Mother Earth." - Robin Wall Kimmerer, Braiding Sweetgrass (photo credit: www.eol.org)

“White man’s footstep, generous and healing, grows with its leaves so close to the ground that each step is a greeting to Mother Earth.” – Robin Wall Kimmerer, Braiding Sweetgrass (photo credit: www.eol.org)

Other Ethnobotany Posts on Awkward Botany: