CAM photosynthesis

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Drought Tolerant Plants: Ice Plants

Among the various strategies plants have for tolerating drought, succulence is easily one of the most common and most successful. A recent article in the new open source journal, Plants People Planet, explores the world of succulent plants, commenting on, among other things, their evolution and extent. At least 83 plant families contain succulent species, and as many as 3-5% of flowering plants are considered succulents.

Succulence involves the storage of water in the cells of one or more plant organs (i.e. roots, stems, or leaves) as a mechanism for surviving drought. One way that succulent species differ is the location and nature of this storage. Some succulents are all cell succulents, meaning that the cells involved in storing water are also involved in carrying out photosynthesis. Other succulents are storage succulents. They have specific cells called hydrenchyma designed for storing water. These cells are non-photosynthetic.

Plants in the family Aizoaceae are storage succulents. Commonly known as the ice plant or carpet weed family, this family consists of hundreds of species and is mainly distributed throughout a region of South Africa known as Succulent Karoo. Species in this family earn the name ice plant thanks to numerous bladder-like cells or hairs that cover their leaves and stems causing them to sparkle or glimmer in the light. Aizoaceae diversity is incredible, and while this post focuses mainly on a few select species, it’s worth browsing through the profiles listed on World of Succulents to appreciate the breadth of forms these plants can take.

common ice plant (Mesembryanthemum crystallinum)

Among many interesting features that plants in this family possess, one particularly fun thing to note is that their flowers, which are unapologetically showy, lack true petals. Instead, what appear as a series of flat, thin petals encircling the center of the flower are actually modified stamens. They act as petals – drawing in pollinators with their bright colors – so calling them petals is acceptable, just not entirely accurate. Another fun fact is that seed pods of plants in Aizoaceae are often hygrochastic – upon getting wet they burst open and expel their seeds.

The photosynthetic pathway in succulents is generally different compared to other plants. Instead of the common C3 pathway, succulents use a pathway called CAM, or Crassulacean Acid Metabolism. CAM photosynthesis is similar to C4 photosynthesis – another photosynthetic pathway common among drought tolerant plants – in that it uses PEP carboxylase instead of rubisco to fix carbon and then sends it to a separate cell to be converted into sugars. In C4 photosynthesis, this whole process happens during the day. CAM photosynthesis differs in that it fixes carbon during the night and then sends it to another cell to be converted into sugars during the day. Fixing carbon at night is a way to avoid the water loss that occurs when collecting carbon dioxide during the daytime.

In discussing Aizoaceae, this is an important consideration because, unlike many other succulents, plants in this family don’t rely solely on CAM photosynthesis, but can instead switch back and forth between C3 and CAM. The ability to do this is likely because they are storage succulents rather than all cell succulents, and because they can do this, they are very efficient carbon fixers.

flowers fading on purple ice plant (Delosperma cooperi)

I live in a region where winter temperatures can dip into the single digits (°F) and sometimes lower,  so my familiarity with ice plants is with cold hardy species and cultivars of the genus Delosperma. If you are familiar with this group of plants, it is most likely thanks to the Plant Select program based in Colorado, particularly the work of Mr. Delosperma himself, Panayoti Kelaidis. Several Delosperma species are cold hardy in the Intermountain West. Thanks to their promiscuous nature, numerous crosses have occurred between species and varieties, resulting in a wide array of flower colors. And speaking of their flowers, the glistening leaves of Delosperma have nothing on their shimmering flowers, some of which may have the ability to temporarily blind you if you’re not careful. Sun is essential though, as they usually close up when shaded.

The cold hardy ice plants of the Delosperma genus are all groundcovers, maintaining a low and creeping profile. Some creep further than others. They are generally not fond of heavy clay soils, and instead prefer soil with good drainage. During the hot, dry days of summer, they appreciate a little water now and then, but watering should be cut off at the end of summer so that they aren’t sitting in saturated soils as winter approaches. They love the sun and will generally flower from late spring throughout the summer. Of course, thanks to their interesting foliage, they catch the eye and provide interest in the garden even when they aren’t flowering.

Fire Spinner® ice plant (Delosperma ‘P001S’)

Within Aizoaceae there are several species that go by the name ice plant that are not so cold hardy. Some are grown as house plants, while others are common in gardens. Still others, like Carpobrotus edulis, were once employed by land managers in California to help control erosion. However, like a number of species introduced for this purpose, C. edulis (commonly known as highway ice plant or hottentot fig) has made itself at home in areas where it wasn’t invited. It has become particularly problematic in coastal ecosystems, spreading quickly across sandy soils and outcompeting native plants. Despite being brought in to control erosion, it actually causes erosion in steep, sandy areas when its carpet-like growth becomes heavy with water and begins sliding down the hill.

highway ice plant (Carpbrotus edulis) carpeting a slope near San Diego – photo credit: Sierra Laverty

Introducing plants to our gardens that come to us from the other side of the globe should be done with caution and care. We don’t want to be responsible for the next invasive species. Since ice plant species have become problematic in California, should we be concerned about cold hardy delospermas? In trialing their plants, invasive qualities are among those that the Plant Select program watches out for, and delospermas seem pretty safe. However, as Kelaidis observes in a blog post from 2014, we should remain vigilant.

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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.