Year of Pollination: Figs and Fig Wasps

This post originally appeared on Awkward Botany in November 2013. I’m reposting an updated version for the Year of Pollination series because it describes a very unique and incredibly interesting interaction between plant and pollinator. 

Ficus is a genus of plants in the family Moraceae that consists of trees, shrubs, and vines. Plants in this genus are commonly referred to as figs, and there are nearly 850 described species of them. The majority of fig species are found in tropical regions, however several occur in temperate regions as well. The domesticated fig (Ficus carica), also known as common fig, is widely cultivated throughout the world for its fruit.

common fig

Common Fig (Ficus carica) – photo credit: wikimedia commons

The fruit of figs, also called a fig, is considered a multiple fruit because it is formed from a cluster of flowers. A small fruit develops from each flower in the cluster, but they all grow together to form what appears to be a single fruit. The story becomes bizarre when you consider the location of the fig flowers. They are contained inside a structure called a syconium, which is essentially a modified fleshy stem. The syconium looks like an immature fig. Because they are completely enclosed inside syconia, the flowers are not visible from the outside, yet they must be pollinated in order to produce seeds and mature fruits.

This is where the fig wasps come in. “Fig wasp” is a term that refers to all species of chalcid wasps that breed exclusively inside of figs. Fig wasps are in the order Hymenoptera (superfamily Chalcidoidea) and represent at least five families of insects. Figs and fig wasps have coevolved over tens of millions of years, meaning that each species of fig could potentially have a specific species of fig wasp with which it has developed a mutualistic relationship. However, pollinator host sharing and host switching occurs frequently.

Fig wasps are tiny, mere millimeters in length, so they are not the same sort of wasps that you’ll find buzzing around you during your summer picnic. Fig wasps have to be small though, because in order to pollinate fig flowers they must find their way into a fig. Fortunately, there is a small opening at the base of the fig called an ostiole that has been adapted just for them.

What follows is a very basic description of the interaction between fig and fig wasp; due to the incredible diversity of figs and fig wasps, the specifics of the interactions are equally diverse.

First, a female wasp carrying the pollen of a fig from which she has recently emerged discovers a syconium that is ready to be pollinated. She finds the ostiole and begins to enter. She is tiny, but so is the opening, and so her wings, antennae, and/or legs can be ripped off in the process. No worries though, since she won’t be needing them anymore. Inside the syconium, she begins to lay her eggs inside the flowers. In doing so, the pollen she is carrying is rubbed off onto the stigmas of the flowers. After all her eggs are laid, the female wasp dies. The fig wasp larvae develop inside galls in the ovaries of the fig flowers, and they emerge from the galls once they have matured into adults. The adult males mate with the females and then begin the arduous task of chewing through the wall of the fig in order to let the females out. After completing this task, they die. The females then leave the figs, bringing pollen with them, and search for a fig of their own to enter and lay eggs. And the cycle continues.

But there is so much more to the story. For example, there are non-pollinating fig wasps that breed inside of figs but do not assist in pollination – freeloaders essentially. The story also differs if the species is monoecious (male and female flowers on the same plant) compared to dioecious (male and female flowers on different plants). It’s too much to cover here, but is a great resource for fig and fig wasp information. Also check out the PBS documentary, The Queen of Trees.



Drought Tolerant Plants: Water Efficient Garden at Idaho State Capitol Building

water efficient garden sign

As drought and threats of drought continue in the western half of the United States, as well as in many other parts of the world, people are increasingly looking for ways to use less water in their landscapes. For many it is a change they are reluctant to make, worried that they will have to sacrifice lush and colorful yards and gardens for drab, dry, gray, and seemingly lifeless ones. Not so, though. The palette of plants that can survive in low water environments is actually quite diverse and contains numerous plants that are just as lush and colorful as some water hogging ones. If planned, planted, and maintained well, a water efficient garden can be incredibly attractive and can even consist of some plants that are comparatively more heavy water users. So, for those who are apprehensive about getting down with brown, don’t fret – there is a better way.

How does one go about creating such a garden? The answer to that is a book on its own – much too long for a single blog post. It also depends who is asking the question, or more specifically, where they are asking it from. Luckily, demonstrations of water-wise gardens are becoming more common. These gardens, planted with regionally appropriate plants and showcasing various water-saving techniques, are great places to start when looking for ideas and motivation. Such gardens can be found at public parks, city and state government buildings, botanical gardens, nurseries and nursery centers, and water company offices. If you are looking to transform your landscape into a more water efficient one, seek out a demonstration garden in your area. It’s a great place to start.

There are several such gardens where I live, one of which is the Water Efficient Garden at the Idaho State Capitol Building in Boise, Idaho. This garden began in 2010 as a partnership between United Water Idaho and the Idaho Capitol Commission. Its mission is to introduce visitors to “low-water native and adaptive plants that thrive in Idaho’s climate.” The plants that were selected for the garden are commonly found at local garden centers and nurseries – an important objective when introducing people to water-wise gardening. The ultimate goal of this garden is to “show homeowners that they can maintain attractive landscaping while conserving water.”

I have my criticisms of this garden regarding plant selection, design, etc., but I’ll spare you those details. I also don’t know the specifics about how this garden is maintained or how often it is watered. All that aside, I am just happy that it exists, and I encourage you to seek out similar gardens in your area. There are numerous approaches to designing and constructing water efficient gardens – again, a book on its own – but demonstration gardens like this are an excellent place to get ideas and learn what other people in your area are doing to conserve water and create landscapes that better reflect the ecology of your region.

United Water Idaho offers a brief introduction to low water gardening here, as well as a list of plants that are in the capitol building garden here.

Blanket Flower (Gaillardia x grandiflora 'Goblin') Plants in the garden are accompanied by a sign with a number on it. The sign corresponds to the plant list that is provided at the entrances to the garden.

Blanket flower (Gaillardia x grandiflora ‘Goblin’). Plants in the garden are accompanied by a sign with a number on it. The sign corresponds to a plant list that is provided at the entrances to the garden.

Dianthus sp.

Dianthus sp.

Coreopsis sp.

Coreopsis sp.

Geranium sp.

Geranium sp.

Liatris sp.

Liatris sp.

A drift of pearly everlasting (Anaphalis margaritacea)

A drift of pearly everlasting (Anaphalis margaritacea)

Purple coneflower (Echinacea purpurea)

Purple coneflower (Echinacea purpurea)

Yellow ice plant (Delosperma nubiginum)

Yellow ice plant (Delosperma nubiginum)

Other “Drought Tolerant Plants” Posts on Awkward Botany:


Blue Sage

Prickly Pears

Growing and Eating Lettuce in Space

Journeying outside of low-earth orbit and setting up long-term or permanent colonies on other planets or moons is fraught with challenges. One obvious challenge is food production. Regular deliveries from Earth are costly and risky, and freshness isn’t always an option. So, perhaps food can be grown on site? NASA is currently exploring this question by carrying out a series of experiments using an aptly named piece of hardware called Veggie, which was delivered and installed on the International Space Station in the spring of 2014. Experiments began shortly thereafter, and last month NASA astronauts finally got to taste the leaves of their labor for the first time.

Veggie is a plant growth chamber that was developed by Orbital Technologies Corporation. It provides environmental conditions – such as light, temperature, and airflow – that are suitable for plant growth. Accompanying the delivery of Veggie were three sets of planting pillows – specially designed pouches that contain growing media, fertilizer, and seeds. The pillows are placed on rooting mats inside Veggie and watered using a wicking system . Light is delivered by red, blue, and green LEDs. The red and blue wavelengths are necessary for plant growth, and the green wavelength helps the plants look more appealing to the astronauts.

Veggie: an expandable plant growth facility designed for the International Space Station - photo credit: NASA/Bryan Onate

Veggie: an expandable plant growth facility designed for growing plants on the International Space Station – photo credit: NASA/Bryan Onate

Two sets of pillows were seeded with a variety of red romaine lettuce called ‘Outredgeous.’ This particular plant was chosen because it is easy to grow, tastes good, and has high nutritional value. The first lettuce harvest was sent back to earth last October for a food safety analysis. Once it was deemed free of harmful bacteria and safe to eat, the astronauts were cleared to start the second round of red romaine, which they did in early July 2015. The third set of planting pillows contain zinnia seeds, and according to statements made by astronaut Scott Kelly on Twitter, it doesn’t sound like those have been grown yet.

After caring for the second round of lettuce plants for 33 days, it was finally time to taste them. The astronauts first cleaned each leaf with citric acid based sanitizing wipes and then sampled the leaves plain. Next they tried them with a little olive oil and balsamic vinegar. They shared their experience in real time via Twitter, which is documented in this New York Times article. They saved a few leaves for their Russian friends who were out on a spacewalk, and then packaged the rest up to be frozen and sent back to Earth for analysis.

Outredgeous Territorial Seed Company

Lactuca sativa ‘Outredgeous’ – the variety of red romaine lettuce grown and eaten by NASA astronauts on the International Space Station (photo credit: Territorial Seed Company)

This isn’t the first time plants have been grown and eaten in space. Russian cosmonauts grew and consumed mizuna (Japanese mustard) back in 2002 using a plant growth chamber developed in collaboration with a lab at Utah State University. They have also used the growth chamber to grow peas, radishes, and other plants. Read more about these experiments here.

Growing plants in space, apart from providing fresh food, offers psychological benefits. In an otherwise sterile and metallic environment, having something green (or red, in the case of the lettuce) to look at and care for has the potential to lift the moods of crew members aboard the space station. NASA scientist, Dr. Gioia Massa, who is overseeing the project sums it up nicely, “The farther and longer humans go away from Earth, the greater the need to grow plants for food, atmosphere recycling, and psychological benefits. I think that plant systems will become important components of any long-duration exploration scenario.”

Want to learn more? Read about the project here, here, and here. Also watch this video about growing plants in space.


Just for fun, there is a great children’s fiction book involving plants in space called June 29, 1999 by David Wiesner which is definitely worth a look.

More “Plants in Space” Posts on Awkward Botany:

Growing Plants on the Moon

Growing Plants in Outer Space

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.”

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.