Year of Pollination: Stamen Movement in the Flowers of Prickly Pears

Last week I made an effort to convince you to add a prickly pear or two to your water-wise gardens. One standout reason to do this is their strikingly beautiful flowers. Apart from being lovely to look at, many prickly pear flowers have a distinct feature that makes them quite fascinating. A demonstration of this feature can be seen in the following video.

 

Stamen movement in response to touch is a characteristic of many species in the genus Opuntia. It isn’t exclusive to Opuntia, however, and can also be seen in Berberis vulgaris, Portulaca grandiflora, Talinum patens, among others. Knowing this makes me want to touch the stamens of any flower I can find just to see what will happen.

The response of stamens to touch has been known for at least a few centuries, but recent research is helping us gain a better understanding of how and why this phenomenon occurs. In general, this movement is thought to assist in the process of cross-pollination. In some cases it may also aid in self-pollination. Additionally, it can have the effect of protecting pollen and nectar from “robbers” (insects that visit flowers to consume these resources but that do not provide a pollination service). Quite a bit of research has been done on this topic, so to simplify things I will be focusing on a paper published in a 2013 issue of the journal, Flora.

In their paper entitled, Intriguing thigmonastic (sensitive) stamens in the plains prickly pear, Cota-Sanchez, et al. studied the flowers of numerous Opuntia polyacantha individuals found in three populations south of Saskatoon, Saskatchewan, Canada. Their objective was to “build basic knowledge about this rather unique staminal movement in plants and its putative role in pollination.” They did this by conducting two separate studies. The first involved observing flower phenology and flower visitors and determining whether the staminal movement is a nasty (movement in a set direction independent of the external stimulus) or a tropism (movement in the direction of the external stimulus). The second involved using high-powered microscopes to analyze the morphology of the stamens to determine any anatomical traits involved in this movement. While the results of the second study are interesting, for the purposes of this post I have chosen to focus only on the findings of the first study.

An important note about the flowers of O. polyacantha is that they are generally protandrous, meaning that the anthers of a single flower release pollen before the stigmas of that same flower are receptive. This encourages cross-pollination. An individual flower is only in bloom for about 12 hours (sometimes as long as 30 hours), however flowering doesn’t occur all at once. The plants in this study flowered for several weeks (from the second week of June to the middle of July).

To determine whether the staminal movement is a nasty or a tropism, the researchers observed insects visiting the flowers. They also manually stimulated the stamens with various objects including small twigs, pencils, and fingers, touching either the inner sides of the filaments (facing the style) or the outer sides (facing the petals). In every observation, the stamens moved in the same direction, “inwards and towards the central part of the flower.” This “consistent unidirectional movement, independent of the area stimulated” led the researchers to categorize the staminal movement of O. polyacantha as thigmonastic. They also observed that staminal movement slowed as the blooming period of an individual flower was coming to an end – “and finally when all the anthers had dehisced, the anthers rested in a clustered position, marking the end of anthesis.” Furthermore, it was observed that “filaments move relatively faster in sunny, warm conditions as opposed to cloudy, cold and rainy days.”

The researchers went on to discuss unique features of the stamens of O. polyacantha. Specifically, the lower anthers contain significantly more pollen than the upper anthers. When the stamens are stimulated, their movement towards the center of the flower results in the lower anthers becoming hidden below the upper anthers. They also noted that small insects less than 5 millimeters in size did not trigger stamen movement. Further observations of the insect vistors helped explain these phenomena.

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A “broad diversity of insects” was observed visiting the flowers, from a variety of bees (bumblebees, honeybees, sweat bees, and mining bees) to bee flies, beetles, and ants. The large bees  were determined to be the effective pollinators of this species of prickly pear. Their large weight and size allows them to push down through the upper anthers to the more pollen-abundant anthers below. After feeding on pollen and nectar, they climb out from the stamens and up to the stigma where they take off, leaving the flower and depositing pollen as they go. Because the bees are visiting numerous flowers in a single flight and the flowers they visit are protandrous, pollen can be transferred from one flower to another and self-pollination can be avoided.

Beetles were observed to be the most common visitors to the flowers; however, they were not seen making contact with the stigma and instead simply fed on pollen and left. Ants also commonly visit the flowers but largely remain outside of the petals, feeding from “extranuptial nectaries.” In short, beetles and ants are not recognized as reliable pollinators of this plant.

Similar results involving two other Opuntia species were found by Clemens Schlindwein and Dieter Wittmann. You can read about their study here.

There are lots of flower anatomy terms in this post. Refresh your memory by visiting another Awkward Botany post: 14 Botanical Terms for Flower Anatomy.

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Drought Tolerant Plants: Prickly Pears

In the introduction to this series about drought tolerant plants, I defended water efficient gardens by claiming they don’t have to be the “cacti-centric” gardens that many visualize upon hearing terms like “xeriscape,” “water-wise,” and “drought tolerant.” And this is absolutely true. However, that won’t stop me from suggesting that such landscapes include a cactus or two. Despite their menacing and potentially dangerous spines, they are actually quite beautiful, and a cactus in bloom is really a sight to behold. Together with a variety of grasses, herbaceous flowering plants, and shrubs, cactus can add unique forms, textures, and focal points that will enhance the look and function of a water-wise garden. This is why I recommend considering cactus, particularly (as far as this post is concerned) one of the many varieties of prickly pears.

The cactus family (Cactaceae) has a native range that is limited to the Americas. Within that range it is expansive, and cactus species can be found in diverse regions from Canada down to Patagonia. The genus Opuntia (the prickly pears) is the most widespread of any genus in the cactus family consisting of at least 300 species found throughout the Americas. Even a brief investigation into Opuntia will reveal that there is considerable controversy as to how many species there actually are and what to call them. This is partly due to the large ranges that species in this genus can have and the diverse habitats they can be found in within those ranges, resulting in a single species having many forms, varieties, and/or subspecies. Hybridization is also common in this genus where ranges overlap, augmenting the challenge of identification.

Generally, prickly pears have flattened stems with spines and glochids emerging from small bumps called areoles. Their flowers are large, showy and a shade of either yellow, orange, or pink and sometimes white. They form fruits that are either fleshy and juicy with a red or purple hue or hard, dry and a shade of brown or tan. The flattened stems are called pads or cladodes and can be quite large in some species, while diminutive and sometimes rounded in others. Some species are without spines, but all have glochids – tiny, barbed, hair-like structures found in clusters on the stems and fruits. While the spines can be painful when they penetrate skin, the glochids are far more irritating as they easily detach themselves from the plant and work their way into the skin of their victims. The fleshy fruits, called tunas, can be eaten after first taking care to remove the glochid-infested outer layer. The young stems of many species can also be eaten – they are referred to as nopales and are common in Mexican cuisine.

Flowers of Opuntia sp. with bee inside flower on the left

Flowers of Opuntia sp. with bee inside flower on the left

Again speaking generally, prickly pears are very easy to propagate and cultivate. Their two main preferences are full sun and well-drained soil. If you are worried that the soil you are planting them in is going to stay too wet for too long, amend it with some gravel. This is especially important if you live in a climate that receives lots of precipitation or that has cold, wet winters. Once established, prickly pears will move around the garden. If that becomes a problem, expanding plants are easily pruned and traveling plants are easily removed.

I live in a climate that requires the selection of cold hardy prickly pears, so I am taking my specific recommendations from two books: Cacti and Succulents for Cold Climates by Leo J. Chance and Hardy Succulents by Gwen Moore Kelaidis. If you live in a warmer climate, your options will be greater. Still, the options for cold regions are pretty numerous, so for the sake of space I am narrowing my list down to a handful that stand out to me at this particular moment.

Three eastern United States species of prickly pears (O. compressa, O. macrorhiza, and O. humifusa) are, according to Chance, “more capable of dealing with wet and cold conditions than almost any other members of the cactus family.” They still require well-drained soil though. An appealing trait is their large, juicy, red fruits that can add garden interest in late summer and fall. Opuntia engelmannii is another species with the potential to tolerate cold, wet conditions. Its size is appealing to me, with pads that reach a foot wide and plants that grow several feet tall. Chance advises finding “a clone that is known to be cold tolerant” and making some space for it, “as it becomes huge in time.” The most cold tolerant prickly pear may be Opuntia fragilis. It is a diminutive plant with a large native range and a variety of forms, some with rounded pads “shaped like marbles.”

Fruits ("tunas") of Opuntia engelmannii - photo credit: www.eol.org

Fruits (“tunas”) of Opuntia engelmannii – photo credit: www.eol.org

Opuntia fragilis 'Frankfurt' - photo credit: wikimedia commons

Opuntia fragilis ‘Frankfurt’ – photo credit: wikimedia commons

Opuntia polyacantha is a prickly pear native to my home state, Idaho. It is found at high elevations throughout the Intermountain West and is also found on the Great Plains. It has many forms and varieties, and its flowers are various shades of pink or yellow. It is a fast growing species and spreads around easily. Other cold hardy species include Opuntia macrocentra (which has a very attractive yellow flower with a red-orange center), Opuntia erinacea (commonly known as hedgehog prickly pear for its abundant, long spines that can obscure the pads), and Opuntia microdisca (a tiny Argentinian prickly pear with pads that barely reach an inch across but, as Chance says, “works very well in a dry rock garden with other miniatures”).

Pads of Opuntia polyacantha

Pads and spines of Opuntia polyacantha

A post about Opuntia could go on indefinitely due to the sheer number of species and their diverse forms and attributes. This is meant merely to pique your interest. The flowers, if nothing else, should certainly interest you. In her book, Kelaidis calls them “improbably beautiful,” and goes on to say that they are “often papery, always glistening and showy.” Chance likens them to “any fancy rose” because they are “extraordinarily large, brightly colored, [and] eye catching.” Next week, as part of Awkward Botany’s Year of Pollination, I will present another reason to be fascinated with the flowers of Opuntia. For now, I will leave you to ponder this word, “thigmonasty.”

Want to learn more about prickly pears? Check out Opuntia Web.