Botany

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Year of Pollination: Most Effective Pollinator Principle and Beyond, part two

“The most effective pollinator principle implies that floral characteristics often reflect adaptation to the pollinator that transfers the most pollen, through a combination of high rate of visitation to flowers and effective deposition of pollen during each visit.” – Mayfield, et al., Annals of Botany (2001) 88 (4): 591-596

In part one, I reviewed a chapter by Jose M. Gomez and Regino Zamora in the book Plant-Pollinator Interactions: From Specialization to Generalization that argues that the most effective pollinator principle (MEPP) “represents just one of multiple evolutionary solutions.” In part two, I summarize a chapter by Paul A. Aigner in the same book that further explains how floral characteristics can evolve without strictly adhering to the MEPP.

maximilian sunflower
Aigner is interested in how specialization develops in different environments and whether or not flowering plants, having adapted to interact with a limited number of pollinators, experience trade-offs. A trade-off occurs when a species or population adapts to a specific environmental state and, in the process, loses adaptation to another state. Or in other words, a beneficial change in one trait results in the deterioration of another. Trade-offs and specialization are often seen as going hand in hand, but Aigner argues that trade-offs are not always necessary for an organism to evolve towards specialization. Plant-pollinator interactions provide an excellent opportunity to test this.

“Flowers demand study of specialization and diversification,” Aigner writes, not only due to their ubiquity, “but because much of the remarkable diversity seen in these organisms is thought to have evolved in response to a single and conspicuous element of the environment – pollination by animals.” If pollinators have such a strong influence on shaping the appearance of flowers, pollination studies should be rife with evidence for trade-offs, but they are not. Apart from not being well-studied, Aigner has other ideas about why trade-offs are not often observed in this scenario.

Aigner is particularly interested in specialization occuring in fine-grained environments. A course-grained environment is “one in which an organism experiences a single environmental state for all of its life.” Specialization is well understood in this type of environment. A fine-grained environment is “one in which an organism experiences all environmental states within its lifetime,” such as “a flowering plant [being] visited by a succession of animal pollinators.” For specialization to develop in a fine-grained environment, a flowering plant must “evolve adaptations to a particular type of pollinator while other types of pollinators are also present.”

It’s important to note that the specialization that Aigner mainly refers to is phenotypic specialization. That is, a flower’s phenotype [observable features derived from genes + environment] appears to be adapted for pollination by a specific type of pollinator, but in fact may be pollinated by various types of pollinators. In other words, it is phenotypically specialized but ecologically generalized. Aigner uses a theoretical model to show that specialization can develop in a fine-grained environment with and without trade-offs. He also uses his model to demonstrates that a flower’s phenotype does not necessarily result from its most effective pollinator acting as the most important selection agent. Instead, specialization can evolve in response to a less-effective pollinator “when performance gains from adapting to the less-effective pollinator can be had with little loss in the performance contribution of the more effective pollinator.”

Essentially, Aigner’s argument is that the agents that are the most influential in shaping a particular organism are not necessarily the same agents that offer the greatest contribution to that organism’s overall fitness. This statement flies in the face of the MEPP, and Aigner backs up his argument with (among other examples) his studies involving the genus Dudleya.

Dudleya saxosa (panamint liveforever) - photo credit: wikimedia commons

Dudleya saxosa (panamint liveforever) – photo credit: wikimedia commons

Dudleya is ecologically generalized. Pollinators include hummingbirds, bumblebees, solitary bees, bee flies, hover flies, and butterflies. “Some Dudleya species and populations are visited by all of these taxa, whereas others seem to be visited by only a subset.” Aigner was curious to see if certain species or populations were experiencing trade-offs by adapting to a particular category of pollinators. Aigner found variations in flower characteristics among species and populations as well as differences in pollinator assemblages that visited the various groups of flowers over time but could not conclude that there were trade-offs “in pollination performance.”

In one study, he looked at pollination services provided by hummingbirds vs. bumblebees as corolla flare changed in size. In male flowers, bumblebees were efficient at removing pollen regardless of corolla flare size, while hummingbirds removed pollen more effectively as corolla flare decreased. Both groups deposited pollen more effectively as corolla flare decreased, but hummingbirds more strongly so. Ultimately, Aigner concluded that “the interactions did not take the form of trade-offs,” or, as stated in the abstract of the study, ” phenotypic specialization [for pollination by hummingbirds] might evolve without trading-off the effectiveness of bumblebees.”

Aigner goes on to explain why floral adaptations may occur without obvious trade-offs. One reason is that different groups of pollinators are acting as selective agents for different floral traits, “so that few functional trade-offs exist with respect to individual traits.” Pollinators have different reasons for visiting flowers and flowers use the pollination services of visitors differently. Another reason involves the “genetic architecture” of the traits being selected for. Results can differ depending on whether or not the genes being influenced are linked to other genes, and genetically based fitness trade-offs may not be observable phenotypically. Further studies involving the genetic architecure of specialized phenotypes are necessary.

And finally, as indicated in part one, pollinators are not the only floral visitors. In the words of Aigner, “if floral larcenists and herbivores select for floral traits in different directions than do pollinators, plants may face direct trade-offs in improving pollination service versus defending against enemies.” These “floral enemies” can have an effect on the visitation rates and per-visit effectiveness of pollinators, which can drastically alter their influence as selective agents.

Like pollination syndromes, the most effective pollinator principle seems to have encouraged and directed a huge amount of research in the field of pollination biology, despite not holding entirely true in the real world. As research continues, a more complete picture will develop. It doesn’t appear that it will conform to an easily digestible principle, but there is no question that, even in its complexity, it will be fascinating.

I will end as I began, with an excerpt from Thor Hanson’s book, The Triumph of Seeds: “The notion of coevolution implies that change in one organism can lead to change in another – if antelope run faster, then cheetahs must run faster still to catch them. Traditional definitions describe the process as a tango between familiar partners, where each step is met by an equal and elegant counter-step. In reality, the dance floor of evolution is usually a lot more crowded. Relationships like those between rodents and seeds [or pollinators and flowers] develop in the midst of something more like a square dance, with couples constantly switching partners in a whir of spins, promenades, and do-si-dos. The end result may appear like quid pro quo, but chances are a lot of other dancers influenced the outcome – leading, following, and stepping on toes along the way.”

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Year of Pollination: Most Effective Pollinator Principle and Beyond, part one

Have you ever considered the diversity of flowers? Why do they come in so many different shapes, sizes, and colors? And why do they produce so many different odors – or none at all? Flowering plants evolved around 140 million years ago, a fairly recent emergence evolutionarily speaking. Along with them evolved numerous species of insects, birds, and mammals. In his book, The Triumph of Seeds, Thor Hanson describes the event this way: “In nature, the flowering plants put sex, seeds, and dispersal on full display, spurring not only their own evolution but also that of the animals and insects with which they became so entwined. In most cases, the diversity of dispersers, consumers, parasites – and, most especially, pollinators – rose right alongside that of the plants they depended upon.”

Speaking of dependence, most flowering plants depend upon pollinators for successful reproduction – it is, for the most part, a mutually beneficial relationship. Even the casual observer of flowers will note that a large portion of the creatures that visit them appear to be pollinators. Thus, it is no wonder that pollination biologists have given pollinators so much credit in shaping the flowers that we see today.

Consider G. Ledyard Stebbins and his Most Effective Pollinator Principle which he defined in a paper published in 1970: “the characteristics of the flower will be molded by those pollinators that visit it most frequently and effectively in the region where it is evolving.” He then goes on to reference pollination syndromes, a phenomenon that describes how the traits of flowers are best suited for their “predominant and most effective vector[s].” In my post about pollination syndromes a few months ago, I discussed how a strict adherence to this concept has waned. In the next two posts, I discuss how the Most Effective Pollinator Principle (MEPP) may not be the best way to explain why flowers look the way they do.

 

To make this argument I am drawing mainly from two chapters in the book Plant-Pollinator Interactions: From Specialization to Generalization. The first is “Ecological Factors That Promote the Evolution of Generalization in Pollination Systems” by Jose M. Gomez and Regino Zamora, and the second is “The Evolution of Specialized Floral Phenotypes in a Fine-grained Pollination Environment” by Paul A. Aigner.

According to Aigner the MEPP “states that a plant should evolve specializations to its most effective pollinators at the expense of less effective ones.” And according to Gomez and Zamora it “states that natural selection should modify plant phenotypes [observable characteristics derived from interactions between a plant’s genes and its surrounding environment] to increase the frequency of interaction [between] plants and the pollinators that confer the best services,” and so “we would expect the flowers of most plants to be visited predominantly by a reduced group of highly effective pollinators.” This is otherwise known as adaptive specialization.

Specialization is something that, in theory, plants are generally expected to evolve towards, particularly in regards to plant-pollinator relationships. Observations, on the other hand, demonstrate the opposite – that specialization is rare and most flowering plants are generalists. However, the authors of both chapters advise that specialization and generalization are extreme ends to a continuum, and that they are comparative terms. One species may be more specialized than another simply because it is visited by a smaller “assemblage” of pollinators. The diversity of pollinators in that assemblage and the pollinator availability in the environment should also be taken into consideration when deciding whether a relationship is specialized or generalized.

That pollinators can be agents in shaping floral forms and that flowering plant species can become specialized in their interactions with pollinators is not the question. There is evidence enough to say that it occurs. However, that the most abundant and/or effective pollinators are the main agents of selection and that specialization is a sort of climax state in the evolutionary process (as the MEPP seems to suggest) is up for debate. Generalization is more common than specialization, despite observations demonstrating that pollinators are drawn to certain floral phenotypes. So, could generalization be seen as an adaptive strategy?

In exploring this question, Gomez and Zamora first consider what it takes for pollinators to act as selective agents. They determine that “pollinators must first benefit plant fitness,” and that when calculating this benefit, the entire life cycle of the plant should be considered, including seed germination rate, seedling survival, fecundity, etc. The ability of a pollinator species to benefit plant fitness depends on its visitation rate and its per-visit effectiveness (how efficiently pollen is transferred) – put simply, a pollinator’s quantity and quality during pollination. There should also be “among-pollinator differences in the evolutionary effect on the plant,” meaning that one species or group of pollinators – through being more abundant, effective, or both – contributes more to plant fitness compared to others. “Natural selection will favor those plant traits that attract the most efficient or abundant pollinators and will also favor the evolution of the phenotypes that cause the most abundant pollinators to also be the most effective.” This process implies possible “trade-offs,” which will be discussed in part two.

When pollinators act as selective agents in this way, the MEPP is supported; however, Gomez and Zamora argue that this scenario “only takes place when some restrictive ecological conditions are met” and that while specialization can be seen as the “outcome of strong pollinator-mediated selection,” generalization can also be “mediated by selection exerted by pollinators…in some ecological scenarios.” This is termed adaptive generalization. In situations where ecological forces constrain the development of specialization and pollinators are not seen as active selection agents, nonadaptive generalization may be occurring.

Gomez and Zamora spend much of their chapter exploring “several causes that would fuel the evolution of generalization” both adaptive and nonadaptive, which are outlined briefly below.

  • Spatiotemporal Variability: Temporal variability describes differences in pollinator assemblages over time, both throughout a single year and over several years. Spatial variability describes differences in pollinator assemblages both among populations where gene flow occurs and within populations. Taken together, such variability can have a measurable effect on the ability of a particular pollinator or group of pollinators to act as a selective agent.
  • Similarity among Pollinators: Different pollinator species can have “equivalent abundance and above all comparable effectiveness” making them “functional equivalents from the plant perspective.” This may be the case with both closely and distantly related species. Additionally, a highly effective pollinator can select for floral traits that attract less effective pollinators.
  • The Real Effects on Plant Fitness: An abundant and efficient pollinator may select for one “fitness component” of a plant, but may “lead to a low overall effect on total fitness.” An example being that “a pollinator may benefit seed production by fertilizing many ovules but reduce seedling survival because it causes the ripening of many low-quality seeds.” This is why “as much of the life cycle as possible” should be considered “in assessing pollinator effectiveness.”
  • Other Flower Visitors: Pollinators are not the only visitors of flowers. Herbivores, nectar robbers, seed predators, etc. may be drawn in by the same floral traits as pollinators, and pollinators may be less attracted to flowers that have been visited by such creatures. “Several plant traits are currently thought to be the evolutionary result of conflicting selection exerted by these two kinds of organisms,” and “adaptations to avoid herbivory can constrain the evolution of plant-pollinator interactions.”

This, of course, only scratches the surface of the argument laid out by Gomez and Zamora. If this sort of thing interests you, I highly encourage you to read their chapter. Next week I will summarize Aigner’s chapter. If you have thoughts on this subject or arguments to make please don’t hesitate to comment or contact me directly. This is a dialogue, dudes.

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Field Trip: Sawtooth Botanical Garden

columbine

It may only be a two and a half hour drive from my house, but until last week I had never visited Sawtooth Botanical Garden in Ketchum, Idaho. The garden is probably not in its prime in the middle of August, but I happened to be in the area so I had to check it out. It’s a small garden – about 5 acres – but I found the space to be well used and full of interesting plants and features. Walking through meandering pathways and around a series of berms, it is easy to get the impression that the garden is larger than it actually is. There were a few areas in obvious need of attention, but as an employee of a non-profit public garden myself, I understand the challenges of maintaining a garden with limited resources. So putting minor issues aside, I thought the garden looked beautiful and I greatly enjoyed my wander through it.

Sawtooth Botanical Garden is in its 11th year. Its mission is to “showcase native and cultivated plants that flourish at high altitude” and to “foster environmental stewardship” of the “region’s unique beauty” by offering “education, events, displays, and plant collections.” Read more about its mission and history here. Brief descriptions of the areas within the garden can also be found on the garden’s website. The interpretive signage describing each area in the garden was well done and one of the highlights of my visit. I didn’t stay long, but I definitely plan on visiting again in the near future. If you ever find yourself in the Wood River Valley, I highly recommend stopping by.

Central area of the garden featuring perennial beds and the Ellen Long Garden Pavillion

Central area of the garden featuring the perennial beds and the Ellen Long Garden Pavillion

Berms in the Alpine Garden with pathway passing through

Berms in the Alpine Garden with pathway passing through

Water feature in the Garden of Infinite Compassion, built in honor of the Dali Lama's visit to the Wood River Valley

Water feature in the Garden of Infinite Compassion, built in honor of the Dalai Lama’s visit to the Wood River Valley several years ago

Alpine strawberry (Fragaria sp.)

Alpine strawberry (Fragaria sp.)

Redtwig dogwood (Cornus stolonifera 'Baileyi')

The fruits of red twig dogwood (Cornus sericea ‘Baileyi’)

cinquefoil

Spring cinquefoil (Potentilla neumanniana)

Spiked speedwell (Veronica spicata 'Red Fox')

Spiked speedwell (Veronica spicata ‘Red Fox’)

Evening primrose (Oenothera sp.)

Evening primrose (Oenothera sp.)

 

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Ethnobotany: White Man’s Foot, part two

Earlier this year, as part of the ethnobotany series, I wrote about plantains (Plantago spp.), of which at least one species is commonly referred to as white man’s foot (or some version of that). Since writing that post, I happened upon a couple of other sources that had interesting and informative things to say about plantains. Rather than go back and update the original post, I decided to make a part two. Hopefully, you find this as interesting as I do. If nothing else, the sources themselves are worth checking out for the additional, fascinating information they contain about all sorts of plants.

plantago_boise capitol building

From The Book of Field and Roadside by John Eastman

Concerning their cosmopolitan nature: “Although both plantains [P. major and P. lanceolata] are Eurasian natives, they have long been thoroughly naturalized global residents; the designation ‘alien’ applies to them in the same sense that all white and black Americans are alien residents.”

In which I learned a new term: “Both species are anthropophilic (associate with humans); they frequent roadsides, parking areas, driveways, and vacant lots, occurring almost everywhere in disturbed ground. Where one species grows, the other can often be found nearby.”

Medicinal and culinary uses according to Eastman: “Plantains have versatile curative as well as culinary properties; nobody need go hungry or untreated for sores where plantains grow. These plants contain an abundance of beta carotene, calcium, potassium, and ascorbic acid. Cure-all claims for common plantain’s beneficial medical uses include a leaf tea for coughs, diarrhea, dysentery, lung and stomach disorders, and the root tea as a mouthwash for toothache. … Their most frequent and demonstrably effective use as a modern herb remedy, however, is as a leaf poultice for insect bites and stings plus other skin irritations. The leaf’s antimicrobial properties reduce inflammation, and its astringent chemistry relieves itching, swelling, and soreness.”

Even the seeds are “therapeutic”: “The gelatinous mucilage surrounding seeds can be readily separated, has been used as a substitute for linseed oil. Its widest usage is in laxative products for providing bulk and soluble fiber called psyllium, mainly derived from the plantain species P. ovata and leafy-stemmed plantain (P. psyllium), both Mediterranean natives.”

Plantain’s “cure-all reputation continues” today: Claims range from a homeopathic cancer remedy to a stop-smoking aid, “supposedly causing tobacco aversion.”

Claims of the healing properties of plantains abound in literature: “John the Baptist, in the lore of the saints, used it as a healing herb; Anglo Saxon gardeners called it the ‘mother of herbs.’ Plantain is ‘in the command of Venus and cures the head by antipathy to Mars,’ according to 17th century English herbalist-astrologist Nicholas Culpeper. Plantains also bear frequent mention in the works of Chaucer and Shakespeare.”

The worst thing plantains have to offer according to Eastman: “the airborne pollen they shed in large amounts, contributing to many hay fever allergies.”

Illustration by Amelia Hansen from The Book of Field and Roadside by John Eastman

Illustration by Amelia Hansen from The Book of Field and Roadside by John Eastman

From Weeds: In Defense of Nature’s Most Unloved Plants by Richard Mabey

Mabey’s too-good-to-paraphrase overview of plantain: “Plantain, ‘the mother or worts,’ is present in almost all the early prescriptions of magical herbs, back as far as the earliest Celtic fire ceremonies. It isn’t clear why such a drab plant – a plain rosette of grey-green leaves topped by a flower spike like a rat’s-tail – should have had pre-eminent status. But its weediness, in the sense of its willingness to tolerate human company, may have had a lot to do with it. The Anglo-Saxon names ‘Waybroad’ or ‘Waybread’ simply mean ‘a broad-leaved herb which grows by the wayside.’ This is plantain’s defining habit and habitat. It thrives on roadways, field-paths, church steps. In the most literal sense it dogs human footsteps. Its tough, elastic leaves, growing flush with the ground, are resilient to treading. You can walk on them, scuff them, even drive over them, and they go on living. They seem to actively prosper from stamping, as more delicate plants around them are crushed. The principles of sympathetic magic, therefore, indicated that plantain would be effective for crushing and tearing injuries. (And so it is, to a certain extent. The leaves contain a high proportion of tannins, which help to close wounds and halt bleeding.)”

On the inclusion of plantains in Midsummer’s Eve rituals: “On Midsummer’s Eve, great bonfires were lit in the countryside, and bundles of wild herbs thrown on them. Most of the plants were agricultural weeds, including St. John’s-wort, corn marigold, corn poppy, mayweed, mugwort, ragwort, plantain, and vervain.”

More about Midsummer’s Eve and the “future-foretelling powers” of this “divination herb, stretching sight into the future”: “On Midsummer’s Eve in Berwickshire, the flowering stems were employed by young women in a charm which would predict whether they would fall in love. It was a delicate, almost erotic process in which the sexual organs of the plantain were used as symbolic indicators. Two of the ‘rat’s-tail’ flowering spikes were picked, and any visible purple anthers removed. The two spikes were wrapped in a dock leaf and placed under a stone. If, by the next day, more anthers had risen erect from the flowering spikes, loves was imminent.”

"Greater - or 'ratstail' - plantain had by this time been nicknamed 'Englishman's foot' by the Native Americans, who had witnessed its prodigious advance in the white man's wake." - Richard Mabey, Weeds: In Defense of Nature's Most Unwanted Plants

“Greater – or ‘ratstail’ – plantain had by this time been nicknamed ‘Englishman’s foot’ by the Native Americans, who had witnessed its prodigious advance in the white man’s wake.” – Richard Mabey, Weeds: In Defense of Nature’s Most Unwanted Plants

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What Shall We Do About Invasive Species?

I think about invasive species a lot. This blog doesn’t really reflect that though. I have been avoiding a deep dive into the subject mainly because there is so much to say about it and I don’t really want this to become “the invasive species blog.” Admittedly, I’m also trying to avoid controversy. Some people have very strong opinions about invasive species, and I don’t always agree. But then an article entitled Taking the long view on the ecological effects of plant invasions appeared in the June 2015 issue of American Journal of Botany. Intrigued by the idea of “taking the long view,” I read the article and decided that now is as good a time as any to start exploring this topic in greater depth.

However, before getting into the article, we should define our terms. “Invasive species” is often used inappropriately to refer to any species that is found outside of its historic native range (i.e. the area in which it evolved to its present form). More appropriate terms for such species are “introduced,” “alien,” “exotic,” “non-native,” and “nonindigenous.” The legal definition of an invasive species (according to the US government) is “an alien species that does or is likely to cause economic or environmental harm or harm to human health.” Even though this definition specifically refers to “alien species,” it is possible for native species to behave invasively.

These terms refer not just to plants but to all living organisms. The term “noxious weed,” on the other hand, is specific to plants. A noxious weed is a plant species that has been designated by a Federal, State, or county government as “injurious to public health, agriculture, recreation, wildlife, or property.” A “weed” is simply a plant that, from a human perspective, is growing in the wrong place, and any plant at any point could be determined to be a weed if a human says so. (I’ll have more to say about human arrogance later in the post.)

Rush skeletonweed (Chondrilla juncea) - labeled a noxious weed in Idaho

Rush skeletonweed (Chondrilla juncea) – labeled a noxious weed in Idaho

The authors of the AJB article (S. Luke Flory and Carla M. D’Antonio) begin by clarifying that “most introduced species are not problematic.” Those that are, however, can “cause significant ecological and economic damage.” This damage is well documented, and it is the reason why billions of dollars are spent every year in the battle against invasive species. But there is a dearth in our research: “less is known about how ecological effects of invasions change over time.” The effects of invasive species could “increase, decrease, or be maintained over decades,” and “multiple community and ecosystem factors” will determine this. For this reason, the authors are calling for “concentrated efforts to quantify the ecological effects of plant invasions over time and the mechanisms that underlie shifting dynamics and impacts.” Armed with this kind of information, managers can better direct their efforts towards invasive species determined to be “the most problematic.”

The authors go on to briefly explain with examples why an invasive species population may decline or be maintained over time, highlighting selected research that demonstrates these phenomena. Research must continue with the aim of improving our understanding of the long term effects of plant invasions. The authors acknowledge that this “will require carefully designed experiments,” “patient and persistent research efforts,” and significant amounts of money. However, they are convinced that through a widespread collaborative effort it can be done. They encourage researchers to deposit data obtained from their research in open source online repositories so that future meta-analyses can be conducted. The information available in these online repositories can be used to develop management plans and help predict “future problematic invasions.”

Considering the amount of time and resources currently spent on confronting invasive species, the approach proposed by the authors of this article is quite reasonable. It seems absurd to continue to battle a problematic species that will ultimately be brought down to more manageable levels by natural causes. It also seems absurd to battle against a species that is essentially here to stay.

Field bindweed (Convulvulus arvensis) - labeled a noxious weed in Idaho

Field bindweed (Convulvulus arvensis) – labeled a noxious weed in Idaho

And that brings me to the point in which I make enemies. Take a look at the terms defined earlier. When we talk about introduced species, we are referring to introductions by humans, whether purposeful or accidental. An “alien” species introduced to a new location by wind, water, or animal (other than human) would be considered a natural introduction, right? If that species becomes established in its new location, it would simply be expanding its range. If a human brought it there, again whether purposefully or accidentally, it would be considered an exotic indefinitely.

Humans have been moving species around since long before we became the humans we are today in the same way that a migratory bird might move a species from one continent to another. At what point during our evolution did our act of moving species around become such a terrible thing?

I will concede that our species has become an incredibly widespread species, able to move about the planet in ways that no other species can. We also have technological advances that no other species comes close to matching. In the time that our species has become truly cosmopolitan, the amount of species introductions that we have participated in has increased exponentially. Leaving ecological destruction in our wake is kind of our modus operandi. I don’t want to make excuses for that, but I also don’t see it unfolding any other way. Give any other species the opportunities we had, and they probably would have proceeded in the same manner. Just consider any of the most notorious invasive species today – “opportunist” is their middle name.

More and more, as we are able to see what we have done, we are making efforts to “fix it.” But how de we rewind time? And if we could, when do we rewind back to? And how do we not “ruin it” again? The earth does not have a set baseline or a condition that it is supposed to be in at any given time. The earth just is. It is operating in a state of randomness, just like everything else in the universe. Any idea of how the earth must look at any given time is purely philosophical – conceived of by humans. I’m not saying that we shouldn’t try to repair the damage, but we should acknowledge that the repairs we’re trying to make are largely for the perpetuation of our own species. Yeah, we’ve developed a soft spot for other species along the way (thankfully), but ultimately we’re just trying to maintain. The earth, on the other hand, would be fine without us.

So, what shall we do about invasive species? I’m not entirely sure. The only thing I’m certain of is that I will continue to ruminate on them and potentially bore you with more blog posts in the future. Until next time…

Puncturevine (Tribulus terrestris) - labeled a noxious weed in Idaho

Puncturevine (Tribulus terrestris) – labeled a noxious weed in Idaho

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Botany in Popular Culture: The Sunset Tree by the Mountain Goats

My obsession with plants means that I see botany everywhere – in the music I listen to, the shows I watch, the books I read, whatever. Just a fleeting mention of something plant related in any type of media will catch my attention, no matter how ancillary it is to the major themes. And that is the impetus behind this series of posts about botany in popular culture. Well that and, believe it or not, I do enjoy many non-plant related things, and this gives me an excuse to write about those things on a plant-centric blog.

TheSunsetTreeFrontCover

The Mountain Goats are a folk rock band formed by John Darnielle in 1991. It could be said that John Darnielle is synonymous with the Mountain Goats, as Darnielle is the chief songwriter and at times has been the only member of the band. The Sunset Tree is the Mountain Goats ninth studio album and only the second album featuring songs that are primarily autobiographical. The album that preceded The Sunset Tree, entitled We Shall All Be Healed, was about Darnielle’s teenage years as a methamphetamine user. The Sunset Tree describes growing up with an abusive stepfather. Heavy topics are kind of the Mountain Goats’ thing.

Darnielle’s lyrics are highly poetic and often nebulous – the listener is left to fill in the gaps. Thus, the storytelling in The Sunset Tree isn’t always direct. However, the scene begins to unfold in the second track, “Broom People,” as Darnielle seems to be describing his childhood living conditions: “all sorts of junk in the unattached spare room,” “dishes in the kitchen sink,” “floor two foot high with newspapers,” “white carpet thick with pet hair.” He also sings of “friends who don’t have a clue; well meaning teachers,” and how he would “write down good reasons to freeze to death in [his] spiral ring notebook.”

“Dance Music” reveals more as Darnielle at 5 or 6 years old is getting “indications that there’s something wrong.” As he sits watching TV, his stepfather is yelling at his mom, then “launches a glass across the room, straight at her head, and [Darnielle] dashes upstairs to take cover.” He turns on his “little record player on the floor” and makes a discovery: “so this is what the volume knob is for.”

A similar scene unfolds in “Hast Thou Considered the Tetrapod,” only this time Darnielle is the victim. He arrives home to find his stepfather asleep, so he sneaks up to his room knowing that if he awakes his stepfather, “there will be hell to pay.” But he does wake up, and he bursts into Darnielle’s room to find him sitting with his headphones on oblivious. The beating begins, and Darnielle sings, “then I’m awake and I’m guarding my face / hoping you don’t break my stereo / because it’s the one thing that I couldn’t live without / and so I think about that and then I sorta black out.” Darnielle describes being “held under these smothering waves” by his stepfather’s “strong and thick-veined hand.” But he remains hopeful that eventually – “one of these days” – he will “wriggle up on dry land.”

That sense of hopefulness can be found throughout the album. In “This Year,” Darnielle is a 17 year old longing to break free. The chorus repeats resolutely: “I am gonna make it through this year if it kills me.” In “Up the Wolves,” he assures us, “there’s gonna come a day when you feel better / you’ll rise up free and easy on that day.”

But there is obviously some anger and frustration expressed as well. Later in “Up the Wolves,” Darnielle sings that he’s going to get himself in “fighting trim” and then makes a series of threats: “I’m gonna bribe the officials, I’m gonna kill all the judges, It’s gonna take you people years to recover from all of the damage.” The song “Lion’s teeth” is a revenge fantasy. Darnielle envisions “the king of the jungle asleep in his car,” and since “nobody in this house wants to own up to the truth,” he takes it upon himself to wrestle the beast. He reaches into the lion’s mouth, grabs onto “one long sharp tooth,” and holds on. The chaos that ensues makes him realize he is “gonna regret the day [he] was born,” but since there is no good way to end it, he is determined to “hold on for dear life.”

The mood lightens during the last two tracks of the album. They seem to be about forgiveness, understanding, and letting go. In “Pale Green Things,” Darnielle tells of hearing from his sister that their stepfather had died “at last, at last.” Upon hearing the news, one of the first memories Darnielle has is of he and his stepfather going to a racetrack to watch horses run. In one scene he recalls looking down at the cracked asphalt and “coming up through the cracks, pale green things.”

It’s a poignant ending to an album full of dark memories. It’s also fitting, as it adds to the bits of hope scattered throughout. Seeing plants push up through concrete or sprout up in detritus collected in gutters and corners of rooftops or even just up out of the dirt in the middle of summer when the ground is hot and bone dry, all of these moments are testaments to the tenacity of living things. Life can, rightfully so, be described as fleeting, short, and fragile – easily snuffed out and erased. But the struggle for life is also fierce, enduring, and relentless. Darnielle’s story is one example of that.

sedums in a hole 2

The “pale green things” that Darnielle saw also symbolize the struggles of the little guy, the underdog, the downtrodden – a tiny, fragile plant pushing its way past solid, suffocating asphalt. It’s a common theme in Darnielle’s music – his latest album is called Beat the Champ, for example. His song “Wild Sage” is also a sign of that ongoing theme.

I work with plants all day, and I am continually awed by them. Daily I am stopped in my tracks, practically forced by some plant to admire one or more of the fascinating features it displays. It doesn’t surprise me that Darnielle would use “pale green things” to express hope and resiliency. Every day I find some kind of hope in plants, that whatever tough thing we are going through, we can one day “wriggle up on dry land” – pale green things pushing up through asphalt, wild sage growing in the weeds.

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Poisonous Plants: Castor Bean

A series of posts about poisonous plants should not get too far along without discussing what may be the most poisonous plant in the world – one involved in high and low profile murders and attempted murders, used in suicides and attempted suicides, a cause of numerous accidental deaths and near deaths, developed for use in biological warfare by a number of countries (including the United States), and used in bioterrorism attacks (both historically and presently). Certainly, a plant with a reputation like that is under tight control, right? Not so. Rather, it is widely cultivated and distributed far beyond its native range – grown intentionally and used in the production of a plethora of products. In fact, products derived from this plant may be sitting on a shelf in your house right now.

Ricinus communis, known commonly as castor bean or castor oil plant, is a perennial shrub or small tree in the spurge family (Euphorbiaceae) and the only species in its genus. It is native to eastern Africa and parts of western Asia but has since been spread throughout the world. It has naturalized in tropical and subtropical areas such as Hawaii, southern California, Texas, Florida, and the Atlantic Coast. It is not cold hardy, but is commonly grown as an ornamental annual in cold climates. It is also grown agriculturally in many countries, with India, China, and Mozambique among the top producers.

Silver maple leaf nestled in the center of a castor bean leaf.

Silver maple leaf nestled in the center of a castor bean leaf.

Castor bean has large palmately lobed leaves with margins that are sharply toothed. Leaves are deep green (sometimes tinged with reds or purples) with a red or purple petiole and can reach up to 80 centimeters (more than 30 inches) across. Castor bean can reach a height of 4 meters (more than 12 feet) in a year; in areas where it is a perennial, it can get much taller. Flowers appear in clusters on a large, terminal spike, with male flowers at the bottom and female flowers at the top. All flowers are without petals. Male flowers are yellow-green with cream-colored or yellow stamens. Female flowers have dark red styles and stigmas. The flowers are primarily wind pollinated and occasionally insect pollinated. The fruits are round, spiky capsules that start out green often with a red-purple tinge and mature to a brown color, at which point they dehisce and eject three seeds each. The seeds are large, glossy, bean-like, and black, brown, white, or often a mottled mixture. They have the appearance of an engorged tick. There is a small bump called a caruncle at one end of the seed that attracts ants, recruiting them to aid in seed dispersal.

Female flowers and fruits forming on castor bean.

Female flowers and fruits forming on castor bean.

All parts of the plant are toxic, but the highest concentration of toxic compounds is found in the seeds. The main toxin is ricin, a carbohydrate-binding protein that inhibits protein synthesis. The seeds need to be chewed or crushed in order to release the toxin, so swallowing a seed whole is not likely to result in poisoning. However, if seeds are chewed and consumed, 1-3 of them can kill a child and 2-6 of them can kill an adult. It takes several hours (perhaps several days) before symptoms begin to occur. Symptoms include nausea, vomiting, severe stomach pain, diarrhea, headaches, dizziness, thirst, impaired vision, lethargy, and convulsions, among other things. Symptoms can go on for several days, with death due to kidney failure (or multisystem organ failure) occurring as few as 3 and as many as 12 days later. Death isn’t imminent though, and many people recover after a few days. Taking activated charcoal can help if the ingestion is recent. In any case, consult a doctor or the Poison Control Center for information about treatments.

The seeds of castor bean are occasionally used to make jewelry. This is not recommended. In The North American Guide to Common Poisonous Plants and Mushrooms, the authors warn that “drilling holes in the seeds makes them much more deadly because it exposes the toxin.” Wearing such jewelry can result in skin irritation and worse. The authors go on to say that “more than one parent has allowed their baby to suck on a necklace of castor beans.” I doubt such parents were pleased with the outcome.

castor bean seeds

Castor beans are grown agriculturally for the oil that can be extracted from their seeds. Due to the way its processed, castor oil does not contain ricin. The leftover meal can be fed to animals after it has been detoxified. Castor oil has been used for thousands of years, dating as far back as 5000 BC when Egyptians were using it as a fuel for lamps and a body ointment, among other things. Over the centuries it has had many uses – medicinal, industrial, and otherwise. It makes an excellent lubricant, is used in cosmetics and in the production of biofuel, and has even been used to make ink for typewriters. One of its more popular and conventional uses is as a laxative, and in her book, Wicked Plants, Amy Stewart describes how this trait has been used as a form of torture: “In the 1920’s, Mussolini’s thugs used to round up dissidents and pour castor oil down their throats, inflicting a nasty case of diarrhea on them.”

A couple of years ago, I grew a small stand of castor beans outside my front door. I was impressed by their rapid growth and gigantic leaves. I also enjoyed watching the fruits form. By the end of the summer, they were easily taller than me (> 6 feet). I collected all of the seeds and still have them today. I knew they were poisonous at the time, but after doing the research for this post, I’m a little wary. With a great collection of castor bean seeds comes great responsibility.

The castor beans that once grew outside my front door.

The castor beans that once grew outside my front door.

There is quite a bit of information out there about castor beans and ricin. If you are interested in exploring this topic further, I recommend this free PubMed article, this Wikipedia page about incidents involving ricin, this article in Nature, and this entry in the Global Invasive Species Database. Also check out Chapter 11 (“Death by Umbrella”) in Thor Hanson’s book, The Triumph of Seeds.

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Year of Pollination: Mosquitoes as Pollinators

It is difficult to have positive feelings about mosquitoes, especially during summer months when they are out in droves and our exposed skin – soft, supple, and largely hair-free – is irresistible to them. We are viewed as walking blood meals by female mosquitoes who are simply trying to produce young – to perpetuate their species just like any other species endeavors to do. Unfortunately, we are left with small, annoying bumps in our skin – red, itchy, and painful – risking the possibility that the mosquitoes that just drew our blood may have passed along any number of mosquito-borne diseases, some (such as malaria) that potentially kill millions of people every year. For this, it is okay to hate mosquitoes and to long for the day of their complete eradication from the planet. However, their ecological roles (and yes, they do have some) are also worth considering.

There are more than 3,500 species of mosquito. Luckily, only 200 or so consume human blood. Mosquitoes go back at least 100 million years and have co-evolved with species of plants and animals found in diverse habitats around the world. Adult mosquitoes and their larvae (which live in standing water) provide food for a wide variety of creatures including birds, bats, insects, spiders, fish, frogs, lizards, and salamanders. Mosquito larvae also help break down organic matter in the bodies of water they inhabit. They even play an important role in the food webs found inside the pitchers of northern pitcher plants (Sarracenia spp.). Interestingly enough, Arctic mosquitoes influence the migration patterns of caribou. They emerge in swarms so big and so voracious that they have been said to kill caribou through either blood loss or asphyxiation.

However, blood is not the main food source of mosquitoes; flower nectar is. Males don’t consume blood at all, and females only consume it when they are producing eggs. Any insect that visits flowers for nectar has the potential to unwittingly collect pollen and transfer it to a nearby flower, thereby aiding in pollination. Mosquitoes are no exception. They have been observed acting as pollinators for a handful of species, and could be acting as pollinators for many more.

Bluntleaved orchid (Platanthera obtusata) is pollinated by mosquitoes. phot credit: wikimedia commons

Bluntleaved orchid (Platanthera obtusata) is pollinated by mosquitoes. photo credit: wikimedia commons

The scientific literature describes the pollination by mosquitoes of at least two plant species: Platanthera obtusata (syn. Habenaria obtusata) and Silene otites. P. obtusata – bluntleaved orchid – is found in cold, wet regions in North America and northern Eurasia. It is pollinated by mosquitoes from multiple genera including several species in the genus Aedes. Mosquitoes visit the flowers to feed on the nectar and, subsequently, pollinia (clusters of pollen) become attached to their eyes and are moved from flower to flower. This scenario likely plays out in other species of Arctic orchids as well*.

S. otites – Spanish catchfly – is a European species that is pollinated by mosquitoes and moths. Researches have been studying the floral odors of S. otites that attract mosquitoes, suggesting that determining the compounds involved in these odors “might lead to the development of new means of pest control and mosquito attractants and repellents.”

Northern House Mosquito (Culex pipiens) - one of the species of mosquitoes that has been observed pollinating Silene otitis. photo credit: www.eol.org

Northern House Mosquito (Culex pipiens) – one of the species of mosquitoes that has been observed pollinating Silene otites. photo credit: www.eol.org

Despite the list of functions that mosquitoes serve in their varied habitats, an article that appeared in Nature back in 2010 argues for wiping mosquitoes off the Earth, stating that “the ecological scar left by a missing mosquito would heal quickly as the niche was filled by other organisms.” And even though “thousands of plant species would lose a group of pollinators,” mosquitoes are not important pollinators of the “crops on which humans depend,” nor do they appear to be the sole pollinator of any single plant species [the species mentioned above are pollinated by other insects as well]. Eliminating mosquitoes, however, is more of a pipe dream than a realistic possibility as our “best efforts can’t seriously threaten an insect with few redeeming features.”

*Speaking of orchids and pollination, endless posts could be written about this incredibly fascinating and diverse group of plants and their equally fascinating and complex mechanisms surrounding pollination. There will be more to come on such topics. Meanwhile, it should be noted that orchids are also a notoriously threatened group of plants. To learn more about orchids and orchid conservation in North America, visit North American Orchid Conservation Center.

Read more about mosquito pollination here.

And now for your listening pleasure:

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Weeds and Wildflowers of the Boise Foothills: June 2015

Boise, Idaho is a beautiful city for many reasons. One feature that makes it particularly attractive are the foothills that flank the city from the southeast to the northwest. The foothills are a transition zone to the mountains that lie to the northeast. Large sections of the foothills have been converted to housing, but much of the area remains as wide open space. There are around 150 miles of trails winding through the foothills that can be accessed from the Boise area. These trails are used frequently by hikers, mountain bikers, dog walkers, bird watchers, trail runners, and horseback riders. The foothills, along with so many other nearby attractions, explains why Boise is such an excellent city for those who love outdoor recreation.

boise foothills trail

I feel embarrassed to say that I had not yet made it into the foothills this year until about a couple weeks ago. I had intended to go for more frequent hikes this year, but life has been in the way. What I was especially curious to see was how the plant life in the foothills changes throughout the year. Because Boise is located in a high desert and receives very little precipitation (especially during the summer months), many of the local wildflowers show themselves in the spring when there is moisture in the soil, after which they wither up and go dormant for the rest of the year.

But there is still lots to see in June. However, it should be noted that when you are hiking in the foothills you must develop an appreciation for weeds, as many of the plants you will see are not native to this area and, in many cases, are in much greater abundance than the plants that are. Species brought in from Europe and Asia have become well established in the Boise Foothills, significantly altering the area’s ecology. One of the major changes has been wildfire frequency. Before weeds like cheatgrass – an annual, shallow-rooted grass imported from Europe – became so prolific in the area, fires were rare, slow moving, and isolated. The continuous, quick burning fuel source provided by dead cheatgrass heightens the risk of more frequent, faster moving, widespread fires, especially in the hot, dry summer months. This threatens plant species that are not adapted to frequent fires.

But this post isn’t about the ecology of the foothills. We can save that for another time. For now, I just wanted to share some of the plants I saw – both native and non-native – on my short walk through a very tiny corner of the Boise Foothills earlier this month.

The trail that I hiked is one of several trails in an area of the Boise Foothills called Hulls Gulch Reserve.

The trail that I hiked is one of several trails in an area of the Boise Foothills called Hulls Gulch Reserve.

 

Bachelor's Buttons (Centaurea cyanus) are native to Europe. They are a common cultivated flower and have escaped from yards into the foothills. They are quite attractive and popular among pollinators. Their flowers and stems are edible so perhaps we should all take to eating them.

Bachelor’s buttons (Centaurea cyanus) are native to Europe. They are a common cultivated flower and have escaped from yards into the foothills. They are quite attractive and popular among pollinators. Their flowers and stems are edible, so perhaps we should all take to eating them.

 

Silverleaf phacelia (Phacelia hastate) - a foothills native that is also a pollinator favorite.

Silverleaf phacelia (Phacelia hastata) – a foothills native and a pollinator favorite.

 

Pale evening primrose (Oenothera pallida) - a foothills native pollinated by nocturnal moths.

Pale evening primrose (Oenothera pallida) – a foothills native pollinated by nocturnal moths.

 

Medusahead (Taeniatherum caput-medusa) is an invasive annual grass from Eurasia. It has an ecological impact similar to cheatgrass (Bromus tectorum).

Medusahead (Taeniatherum caput-medusae) is an invasive annual grass from Eurasia. It has an ecological impact similar to cheatgrass (Bromus tectorum).

 

The fruits of nineleaf biscuitroot (Lomatium triternatum), a spring flowering plant in the carrot family (Apiaceae).

The fruits of nineleaf biscuitroot (Lomatium triternatum), a native spring wildflower in the carrot family (Apiaceae).

 

Fruits forming on antelope bitterbrush (Purshia tridentata), one of several shrubs native to the Boise Foothills.

Fruits forming on antelope bitterbrush (Purshia tridentata), one of several shrubs native to the Boise Foothills.

 

Rubber rabbitbrush (Ericameria nauseosa), a native shrub that flowers in late summer.

Rubber rabbitbrush (Ericameria nauseosa), a native shrub that flowers in late summer.

 

Lichens on the branch of basin big sagebrush (Artemisia tirdentata sbsp. tridentata) another common native shrub.

Lichens on the branches of basin big sagebrush (Artemisia tridentata subsp. tridentata), another common native shrub.

 

Tall tumblemustard (Sisymbrium altissimum) an introduced species and one of many tumbleweed species in the western states.

Tall tumblemustard (Sisymbrium altissimum) – an introduced species and one of many tumbleweed species in the western states.

 

Little spider atop the flowers of western yarrow (Achilea millefolium), a foothills native.

A little spider atop flowers of western yarrow (Achilea millefolium var. occidentalis), a foothills native.

Learn more about the Boise Foothills here and here.

Where have you been hiking lately?

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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.

SAMSUNG

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.

Recently I received a note from a reader requesting that I include a link to subscribe to this blog’s RSS Feed. I have now made that available, and it can be found at the top of the sidebar.