Insect Killer, Plant Symbiont

There has been an uptick in conversations about plant-fungal interactions recently. News of trees communicating via a vast subterranean network of fungal threads has everyone looking at forests like one big commune. Though it feels nice to think of these relationships as altruistic, such simplified takes on the subject overlook the fact that plants and the mycorrhizal fungi they partner with have entered into a mutual exchange, allowing each player to gain from the interaction.

The reciprocity of these relationships are exquisitely illustrated in the partnering of fungi in the genus Metarhizium and their botanical hosts. Metarhizium are predominantly insect pathogens, invading the bodies of soil-dwelling insects, killing them, and absorbing nutrients like nitrogen that are locked up within their tissues. Though extremely good at obtaining compounds like nitrogen from insects, these fungi can not readily access the carbon they need to survive. That is where plants come in.

Plants are experts at producing carbon-based compounds. Via photosynthesis, they break apart CO2 molecules and turn them into carbon-rich sugars for food. However, they need nitrogen to do this. Unfortunately for plants, most of the nitrogen on our planet is locked up in forms they can’t readily access. It is likely that plants’ relative inefficiency at obtaining the nitrogen they need to survive is a major driving force for the partnering between plants and soil-dwelling fungi.

A beetle grub infected by a Metarhizium fungus. Photo by CSIRO (CC BY 3.0)

Over the last few years, scientists studying the relationship between Metarhizium and plants have discovered that a fascinating and ecologically important exchange has evolved among these organisms. When plants are presented with adequate nitrogen, many species will end up over-producing carbohydrates. Their fungal partners are the ones to benefit from this as those excess carbohydrates are fed to the fungi living on or in the plants’ roots. Indeed, via some complex experiments using isotopes of carbon and nitrogen, scientists were able to demonstrate that killing and eating insects isn’t the only way Metarhizium fungi make a living.

In addition to eating insects, Metarhizium also form mycorrhizal relationships with the roots of numerous plant species from grasses to beans. In doing so, they are able to obtain carbohydrates. However, the plants aren’t giving their photosynthates away for free. In exchange, the fungi are providing them with ample nitrogen that was obtained by infecting and digesting their insect prey. By tracing the path of carbon and nitrogen isotopes between fungi and plants, scientists found that the fungi were supplying the plants directly with insect-derived nitrogen.

This may not sound terribly surprising. After all, this is more or less how most mycorrhizal interactions work. However, the fact that an insect-killing fungus is transferring nitrogen from insect to plant directly, rather than from already decomposed materials in the soil reveals a rather novel pathway in the nitrogen cycle of our planet. Metarhizium is an extremely common and widespread genus of fungi and it is likely that these relationships are not unique to the plants used in these studies. The wide-spread nature of these relationships means that this way of cycling nitrogen and carbon through an ecosystem is also extremely common and wide spread.

It is important to remember that relationships like these are a benefit to plants and fungi alike (sorry insects). Both parties stand to gain from the mutualism. It isn’t that plants are plugging into this system and using it to help each other out. To me, it makes far more sense that fungi like Metarhizium benefit from keeping as many healthy plants in their network as possible. We can’t forget that like plants, fungi are organisms fighting to survive long enough to get their genes into the next generation. Mutualisms are not altruisms. They are mutual exchanges that benefit both parties.

Photo Credits: [1] [2]

Further Reading: [1] [2] [3]

How radioactive carbon from nuclear bomb tests can tell us what parasitic orchids are eating

Yoania japonica. Photo by Qwert1234 licensed by CC BY-NC-SA 4.0

Historically, non-photosynthetic plants were defined as saprotrophs. It was thought that, like fungi, such plants lived directly off of decaying materials. Advances in our understanding have since revealed that parasitic plants don’t do any of the decaying themselves. Instead, those that aren’t direct parasites on the stems and roots of other plants utilize a fungal intermediary. We call these plants mycoheterotrophs (fungus-eaters). Despite recognition of this strangely fascinating relationship, we still have much to learn about what kinds of fungi these plants parasitize and where most of the nutritional demands are coming from.

It is largely assumed that most mycoheterotrophic plants are parasitic on mycorrhizal fungi. This would make them indirect parasites on other photosynthetic plants. The mycorrhizal fungi partner with photosynthetic plants, exchanging soil nutrients for carbon made by the plant during photosynthesis. However, this is largely assumed rather than tested. New research out of Japan has shown a light on what is going on with some of these parasitic relationships and the results are a bit surprising. What’s more, the methods they used to better understand these parasitic relationships are pretty clever to say the least.

Cyrtosia septentrionalis Photo by Qwert1234 licensed by CC BY-NC-SA 4.0

Photosynthesis involves the uptake of and subsequent breakdown of CO2 from the atmosphere. The carbon from CO2 is then used to build carbohydrates, which form the backbone of most plant tissues. Not all carbon is created equal, however, and by looking at ratios of different carbon isotopes in living tissues, scientists can better understand where the carbon came from. For this research, scientists utilized an isotope of carbon called 14C.

Eulophia zollingeri photo by Vinayraja licensed by CC BY-NC-SA 3.0

14C is special because it is not as common in our atmosphere as other isotopes of carbon such as 12C and 13C. One of the biggest sources of 14C in our atmosphere were nuclear bomb explosions. From the 1950’s until the Partial Nuclear Test Ban in 1963, atomic bomb tests were a regular occurrence. During that time period, the concentration of 14C in our atmosphere greatly increased. Any organism that was fixing carbon into its tissues during that span of time will contain elevated levels of 14C compared to the other carbon isotopes. Alternatively, anything fixing carbon today, say via photosynthesis, will have considerably reduced levels of 14C in its tissues.

Gastrodia elata Photo by Qwert1234 licensed by CC BY-NC-SA 4.0

By looking at the ratios of 14C in the tissues of parasitic plants, scientists reasoned that they could assess the age of the carbon being utilized. If more 14C was present, the source of the carbon could not come from today’s atmosphere and therefore not from recent photosynthesis. Instead, it would have to come from older sources like decaying wood of long-dead trees. In other words, if parasitic plants were high in 14C, then the scientists could reasonably conclude that they were parsitizing wood-decaying saprotrophic fungi. If the plants were high in 12C or 13C, then they could conclude that they were partnering with mycorrhizal fungi instead, which were obtaining carbon from present-day photsynthesis.

The researchers looked at 10 different species of parasitic plants across Japan, most of which were orchids. They analyzed their tissues and ran analyses on the carbon molecules within. What they found is that 6 out of the 10 plants contained much higher levels of 12C and 13C in their tissues, which points to mycorrhizal fungi as their host. However, for the 4 remaining species (Gastrodia elata, Cyrtosia septentrionalis, Yoania japonica and Eulophia zollingeri), the ratios of 14C were considerably higher, meaning their host fungi were eating dead wood, not partnering with photosynthetic plants near by.

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Indeed, it appears that at least some mycoheterotrophic plants are benefiting from saprotrophic rather than mycorrhizal fungi. Those early assumptions into the livelihood of such plants were not as far off the mark after all. This is very exciting research that opens the door to a much deeper understanding of some of the strangest plants on our planet.

LEARN MORE ABOUT MYCOHETEROTROPHIC PLANTS IN EPISODE 234 OF THE IN DEFENSE OF PLANTS PODCAST

Photo Credits: [1] [2] [3] [4] [5]

Further Reading: [1] [2]

The Round Leaved Orchid

In the northern temperate regions of North America, late June marks the beginning of what I like to call orchid season. If you're lucky you may stumble across one of these rare beauties in full bloom. Their diversity in shape and size are mainly a result of the intricate evolutionary relationships they have formed with their pollinators. I spend much of my time botanizing trying to locate and photograph these botanical curiosities and any time I get to meet a new species is a very special time indeed. 

Take the round leaved orchid (Platanthera orbiculata) for example. For years I have only known this species as two round leaves that are slightly reminiscent of the phaleanopsis orchids you see for sale in nurseries and grocery stores. The leaves can be quite large too. With their glossy appearance, they are the easiest way to locate this plant.

When conditions are right and the plants have enough stored energy they will begin to flower. Rising from the middle of the pair of leaves is a decent sized inflorescence loaded with greenish white flowers. The flowers are interesting structures. Not particularly colorful, they have a long white lip and considerable green nectar spurs. There are said to be two varieties of this species, each being characterized by the length of the nectar spur. Unlike many orchids that offer no reward to pollinators, P. orbiculata produces nectar. The flowers are pollinated by noctuid moths, which is probably why they are white in color. Whereas most lepidopteran pollinated orchid attach their pollinia to the proboscis of the butterfly or moth, P. orbiculata attaches its pollinia to the eyes of visiting moths. 

If this isn't strange enough, the pollinia themselves have some of their own intriguing adaptations. Visiting moths take a certain amount of time to successfully access the nectar from the nectar spur. If the plant is to avoid wasting precious pollen on itself, then it must find a way to delay this process. The pollinia are the solution to this. When first attached to the eyes, the pollinia stick straight up. This keeps them away from the female parts of the plant as the moth feeds. Only after enough time has elapsed will the stalks of the pollinia begin to bend forward. At this point the moth will hopefully have moved on to the flowers of an unrelated individual. Pointing straight forward, they are now perfectly positioned to transfer pollen. 

Like all orchids, P. orbiculata relies on specialized mycorrhizal fungi for germination and survival. At the beginning of its life, P. orbiculata relies solely on the fungi for sustenance. Once it has enough energy to produce leaves it will repay the fungi by providing carbohydrates. However, the relationship is not over at this point. Every spring, P. orbiculata produces a new set of leaves as well as a whole new root system. The fungi supply a lot of energy for this process and if the plant is disturbed (ie. dug up by greedy poachers) or browsed upon, it is likely that it will not recover from the stress and it will die. The mycorrhizal fungi it relies on live on rotting wood so finding well rotted logs is a good place to start searching for this species. With declining populations throughout much of its range, it is important to remember to enjoy it where it grows. Leave wild orchids in the wild!

Further Reading: [1] [2] [3] [4]

Crab Spiders and Pitcher Plants: A Dynamic Duo

Most pitcher plants in the genus Nepenthes seem pretty adept at catching prey. These plants specialize in nutrient-poor soils and their carnivorous habit evolved as a means of supplementing their nutritional needs. Despite the highly evolved nature of their pitfall traps (which are actually modified leaves), Nepenthes aren’t perfect killing machines. In fact, some get a helping hand from seemingly unlikely partners.

Spend enough time reading about Nepenthes in the wild and you will see countless mentions of arthropods hanging around their pitchers. Some of these inevitably become prey, however, there are others that appear to be taking advantage of the plant. Nepenthes don’t passively trap arthropods. Instead, they lure them in with bright colors and the promise of tasty treats like nectar. This is not lost on predators like spiders, who are frequent denizens of pitcher mouths.

Most notable to Nepenthes specialists are some of the crab spiders that frequently haunt Nepenthes traps. These wonderful arachnids sit at the mouth of the pitcher and ambush any insects that try to pay it a visit. Often times both predator and prey fall down into the pitcher, however, thanks to a strand of silk, the spiders easily climb back out with their meal. This may seem like bad news for the pitcher, however, recent research based out of the National University of Singapore has shown that this relationship is not entirely one sided.

By studying the interactions between spiders and pitcher plants both in the lab and in the field, ecologists discovered that at least one species of pitcher plant (Nepenthes gracilis) appears to benefit greatly from the presence of crab spiders. The key to understanding this relationship lies in the types of prey N. gracilis is able to capture when crab spiders are and are not present.

Not only did the presence of a resident crab spider increase the amount of prey in each Nepenthes pitcher, it also changed the types of insects that were being captured. Crab spiders are ambush predators that frequently attack prey much larger than themselves. It may seem as if this is a form of food robbery on the part of the crab spider but the spiders can’t eat everything. When they have eaten their fill, the spiders discard the carcass into the pitcher where the plant can make quick work digesting it for its own benefit.

Over time, simply having a spider hunting on the trap led to a marked increase in the number of insects in each pitcher compared to those without a spider. Even if these meals are already half eaten, the plant still gains nutrients. Additionally, the types of prey captured by pitchers with and without crab spiders changed. The spiders were able to capture and subdue insects like flesh flies, which normally aren’t captured by Nepenthes pitchers. As such, the resident crab spiders make available a larger suite of potential prey than would be available if they weren’t using the pitchers as hunting grounds.

The crab spiders may benefit the pitcher plant in other ways as well. Research on crab spiders has shown that their bodies are covered in pigments that register high in the UV spectrum. Insects can see UV light and often use it as a means of finding flowers as plants often produce UV-specific pigments in their floral tissues. The wide array of UV patterns on flowers are there to guide their pollinators into position. Researchers have documented that insects are actually more likely to visit flowers with crab spiders than those without, which has led to the idea that UV pigments in crab spiders actually act as insect attractants. Visiting insects simply cannot resist the UV stimulus and quickly fall victim to the resident crab spider.

Could it be that by taking up residence on a Nepenthes pitcher, the crab spiders are increasing the likelihood of insects visiting the traps? This remains to be seen as such questions did not fall under the scope of this investigation. That being said, it certainly offers tantalizing evidence that there is more to the Nepenthes-crab spider relationship. More work is needed to say for sure but the closer we look at such interactions, the more spectacular they become!

Photo Credits: [1] [2] [3] [4]

Further Reading: [1] [2] [3]

An Introduction to Hornworts

Anthoceros sp. Photo by Bramadi Arya licensed under CC BY-SA 4.0

When was the last time you thought about hornworts? Have you ever thought about hornworts? If you answered no, you aren’t alone. Despite their global distribution, these tiny plants receive hardly any attention and that is a shame. Hornworts (Anthocerotophyta) have been around for a very long time. In fact, it is likely that they were some of the first plants to colonize the land roughly 300 - 400 million years ago.

To be fair, hornworts aren’t known for their size. They are generally small plants, though their colonies can form impressive mats. To find them, one must try looking in and among rocks, bare patches of soil, or pretty much anywhere enough moisture builds up to supply their needs. They tend to enjoy nutrient-poor substrates but I would hesitate to say that with any certainty. No matter where you live, from the tundra to the tropics, there is probably a hornwort native to your neck of the woods.

Dendroceros sp. Photo by J.Ziffer licensed under public domain

How many different species of hornwort there are is apparently the subject of some debate. Some authors recognize upwards of 300 species whereas others suggest the real number hangs somewhere around 150. Regardless of the exact numbers, hornworts belong to one of six genera: Anthoceros, Dendroceros, Folioceros, Megaceros, Notothylas and Phaeoceros. Fun fact, the suffix ‘ceros’ at the end of each genus is derived from the Latin word for ‘horn.’

The reason they are called hornworts is because of their reproductive structures or “sporophytes.” Similar to their moss and liverwort cousins, hornworts undergo an alternation of generations in order to reproduce sexually. The green gametophytes house the sexual organs - antheridia if they are male and archegonia if they are female. After fertilization, a sporophyte begins to grow, which will go on to produce and disseminate spores. However, the way in which the hornwort sporophyte forms is a bit different from what we see in mosses and liverworts.

Alternation of generations in hornworts. Photo by Mariana Ruiz (LadyofHats) licensed under public domain

Upon fertilization, the zygote begins to divide into a bulbous mass of cells affectionately referred to as "the foot.” This foot remains within the gametophyte throughout the lifetime of the hornwort, depending on the gametophyte for water and nutrients. Even more peculiar is the the fact that the growing point of the sporophyte is at the base rather than the tip. As such, the horn of each hornwort could continue to grow upwards until it is damaged in some way.

The horn itself is an amazing structure. Whereas the outside layers of tissue are merely structural, the internal tissues differentiate into two different types - spores and pseudo-elaters. Pseudo-elaters expand and contract as humidity fluctuates so as the sporophyte splits to release the spores, the pseudo-elaters dehydrate and snap like tiny spore catapults, thus aiding in their dispersal.

Megaceros flagellaris. Photo by Dr. Scott Zona licensed under CC BY-NC 2.0

Of course, reproduction is the main goal but to get to that point, hornworts must grow and mature. How they manage to survive is incredible because it is a reminder that what are often thought of as “primitive” plants are actually far more advanced than we give them credit for. The main body of the hornwort gametophyte is a thin layer of cells that spread out to form a tiny, green mat. This is the structure you are most likely to encounter.

Inside each cell is a single chloroplast. In most hornworts, the chloroplast does not exist in isolation. Instead, it is fused with other organelles into a structure called a “pyrenoid.” The pyrenoid functions as both a center for photosynthesis and a food storage organ. This is unique as it relates to terrestrial plants but quite common in algae. Another odd fact about hornwort anatomy are the presence of tiny cavities scattered throughout their tissues. These cavities form as clusters of hornwort cells die. They then fill with a special mucilage that appears to invite colonization by nitrogen-fixing cyanobacteria. The cyanobacteria set up shop within the cavities and provides the hornwort with supplemental nitrogen in return for a place to live.

Anthoceros agrestis photo by BerndH licensed under CC BY-SA 3.0

Cyanobacteria aren’t the only organisms to have partnered with hornworts either. Mycorrhizal fungi also enter into the picture. A study done back in 2013 actually found that a wide variety of fungi will partner with hornworts which suggests that this symbiotic relationship is much more ancient and versatile than we once thought. Fungi cluster around parts of the gametophyte that produce root-like structures called “rhizoids,” offering nutrients in return for carbohydrates.

All in all, I think it is safe to say that hornworts are remarkable little plants. Though they can sometimes be difficult to find and properly identify, they nonetheless offer plenty of inspiration for the botanically inclined mind. We can all do better by tiny plants like the hornworts. They have been on land for an incredible amount of time and they definitely deserve our respect and admiration.

Photo Credits: [1] [2] [3] [4] [5]

Further Reading: [1] [2] [3]

Life With Endophytic Fungi

Endophytic fungi living in the cells of a grass leaf. Photo by Nick Hill (Public Domain)

Talk about plants long enough and fungi eventually make their way into the conversation. These two walks of life are inextricably linked and probably have been since the earliest days. At this point we are well aware of beneficial fungal partners like mycorrhizae or pathogens like the cedar apple rust. Another type of relationship we are only starting to fully appreciate is that of plants and endophytic fungi living in their above ground tissues. 

Endophytic fungi have been discovered in many different types of plants, however, it is best studied in grasses. The closer we look at these symbiotic relationships, the more complex the picture becomes. There are many ways in which plants can benefit from the presence of these fungi in their tissues and it appears that some plants even stock their seeds with fungi, which appears to give their offspring a better chance at establishment. 

To start, the benefits to the fungi are rather straight forward. They get a relatively safe place to live within the tissues of a plant. They also gain access to all of the carbohydrates the plants produce via photosynthesis. This is not unlike what we see with mycorrhizae. But what about the plants? What could they gain from letting fungi live in and around their cells?

One amazing benefit endophytic fungi offer plants is protection. Fungi are famous for the chemical cocktails they produce and many of these can harm animals. Such benefits vary from plant to plant and fungi to fungi, however, the overall effect is largely the same. Herbivores feeding on plants like grasses that have been infected with endophytic fungi are deterred from doing so either because the fungi make the plant distasteful or downright toxic. It isn't just big herbivores that are deterred either. Evidence has shown that insects are also affected.

There is even some evidence to suggest that these anti-herbivore compounds might have influences farther up the food chain. It usually takes a lot of toxins to bring down a large herbivore, however, some of these toxins have the potential to build up in the tissues of some herbivores and therefore may influence their appeal to predators. Some have hypothesized that the endophytic fungal toxins may make herbivores more susceptible to predators. Perhaps the toxins make the herbivores less cautious or slow them down, making them more likely targets. Certainly more work is needed before anyone can say for sure.

Italian ryegrass (Lolium multiflorum) is one of the most studied grasses that host endophytic fungi. Photo by Matt Lavin licensed under CC BY-SA 2.0

Another amazing example deals with parasitoids like wasps that lay their eggs in other insects. Researchers found that female parasitoid wasps can discriminate between aphids that have been feeding on plants with endophytic fungi and those without endophytic fungi. Wasp larvae developed more slowly and had a shorter lifespan when raised in aphids that have fed on endophytic fungi plants. As such, the distribution of plants with endophytic symbionts may have serious ramifications for parasitoid abundance in any given habitat.

Another benefit these endophytic fungi offer plants is increased photosynthesis. Amazingly, some grasses appear to photosynthesize better with endophytic fungi living in their tissues than plants without fungi. There are many mechanisms by which this may work but to simplify the matter, it appears that by producing defense compounds, endophytic fungi allow the plant to redistribute their metabolic processes towards photosynthesis and growth. In return, the plants produce more carbohydrates that then feed the fungi living in their tissues. 

One of the most remarkable aspects about the relationship between endophytic fungi and plants is that the plants can pass these fungi on to their offspring. Fungi are able to infect the tissues of the host plants' seeds and therefore can be carried with the seeds wherever they go. As the seedlings grow, so do the fungi. Some evidence suggests this gives infected seedlings a leg up on the competition. Other studies have not found such pronounced effects.

Still other studies have shown that it may not be fungi in the seeds that make a big difference but rather the fungi present in the decaying tissues of plants growing around them. Endophytic fungi have been shown to produce allelopathic compounds that poison neighboring plants. Areas receiving lots of plant litter containing endophytic fungi produced fewer plants but these plants grew larger than areas without endophytic fungi litter. Perhaps this reduces competition in favor of plant species than can host said endophytes. Again, this has potentially huge ramifications for the diversity and abundance of plant species living in a given area.

We are only beginning to understand the role of endophytic fungi in the lives of plants and the communities they make up. To date, it would appear that endophytic fungi are potentially having huge impacts on ecosystems around the globe. It goes without saying that more research is needed.

Photo Credits: [1] [2]

Further Reading: [1] [2] [3] [4] [5] [6]

The Bladderwort Microbiome Revealed

Photo by Stefan.lefnaer licensed under CC BY-SA 3.0

The bladderworts (Utricularia spp.) are among the most cosmopolitan groups of carnivorous plants on this planet. Despite their popularity, their carnivorous habits have been subject to some debate. Close observation reveals that prey capture rates are surprisingly low for most species. This has led some to suggest that the bladderworts may be benefiting from more passive forms of nutrient acquisition. To better understand how these plants utilize their traps, a team of researchers decided to take a closer look at the microbiome living within. 

The team analyzed the trap fluid of a handful of floating aquatic bladderwort species - U. vulgaris, U. australis, and U reflexa. In doing so, they uncovered a bewildering variety of microorganisms perfectly at home within the bladderwort traps. Thanks to sophisticated genetic tools, they were able to classify these microbes in order to investigate what exactly they might be doing inside the traps. 

Their findings were quite astonishing to say the least. The traps of these plants harbor extremely rich microbial communities, far richer than the microbial diversity of other carnivorous plant traps. In fact, the richness of these microbial communities were more akin to the richness seen in the rooting zone of terrestrial plants or the gut of a cow. In terms of the species present, the microbial communities of bladderwort traps most closely resembled that of the pitchers of Sarracenia species as well as the guts of herbivorous iguanas.

Photo by Michal Rubeš - http://www.darwiniana.cz/vamr/?page=obrazek&id=914 licensed under CC BY 4.0

The similarities with herbivore guts is quite remarkable. Its not just coincidental either. The types of microbes they found weren't new to science but their function was a bit of a surprise. A large percentage of the bacteria living within the fluid are famously known for producing enzymes that digest complex plant tissues. Similarly, the team found related microbe groups that specialize on anaerobic fermentation. These types of microbes in particular are largely responsible for the breakdown of plant materials in the rumen of cattle.

As it turns out, the microbes living within the traps of these bladderworts are serving a very important purpose for the plant - they are breaking down plant and algae cells that find their way into the traps each time they open and close. In doing so, they give off valuable nutrients that the bladderworts can then absorb and utilize. Let me say that again, the bacteria living in bladderwort traps are digesting algae and other plant materials that these carnivorous plants can then absorb.

Now these bacteria are also responsible for producing a lot of methane in the process. Interestingly enough, the team was not able to detect measurable levels of methane leaving the traps. This would be odd if it wasn't for the community of methane-feeding microbes also discovered living within the traps. The team believes that these organisms metabolize all of the methane being produced before it can escape the traps. 

As remarkable as these findings are, I don't want to give the impression that these carnivorous plants have taken up a strict vegetarian lifestyle. The team also found myriad other microorganisms within the bladder traps, many of them being carnivores themselves. The team also found a rich protist community. A majority of these were euglenids and ciliates. 

These sorts of protists are important microbial predators and the numbers recorded within the traps suggest that they are a rather significant component of these trap communities. As they chase down and consume bacteria and other protists, they release valuable nutrients that the plants can absorb and utilize. Numbers of these predatory protists were much higher in older traps, which have had much more time to accumulate a diverse microbiome. Astonishingly, it is estimated that the protist communities can cycle the entire contents of the bladderwort traps upwards of 4 or 5 times in a 24 hour period. That is some serious turnover of nutrients!

The protists weren't the only predators found within the traps either. There are also a considerable amount of bacterial predators living there as well. These not only cycle nutrients in similar ways to the protist community, it is likely they also exhibit strong controls on the biodiversity within this miniature ecosystem. In other words, they are considered keystone predators of these microcosms.

Also present within the traps were large amounts of fungal DNA. None of the species they found are thought to actually live within the traps. Rather, it is thought that they are taken up as spores blown in from the surrounding environment. Exactly how these organisms find themselves living inside bladderwort traps is something worth considering. The plants themselves are known for being covered in biomfilms. It is likely that many of the organisms living within the traps were those found living on the plants originally. 

Taken together, the remarkable discovery of such complex microbial communities living on and within these carnivorous plants shows just how complex the ecology of such systems really are. Far from the active predators we like to think of them as, the bladderworts nonetheless rely on a mixture of symbiotic orgnaisms to provide them with the nutrients that they need. The fact that these plants are in large part digesting plant and algae materials is what I find most astonishing.

Essentially, one can almost think of bladderworts as plants adorned with tiny, complex cow stomachs, each utilizing their microbial community to gain as much nutrients as they can from their living environment. The bladderworts gain access to nutrients and the microbes get a place to live. The bladderworts really do seem to be cultivating a favorable habitat for these organisms as well. Analysis of the bladder fluid demonstrated that the plants actively regulate the pH of the fluid to maintain their living community of digestive assistants. In doing so, they are able to offset the relative rarity of prey capture. Keep in mind that this research was performed on only three species of bladderwort originating from similar habitats. Imagine what we will find in the traps of the multitude of other Utricularia species.

Photo Credits: [1] [2] [3]

Further Reading: [1]

The Nitrogen-Fixing Abilities of Cycads

Photo by Daderot Public Domain

Long before the first legumes came onto the scene, the early ancestors of Cycads were hard at work fixing atmospheric nitrogen. However, they don't do this on their own. Despite being plentiful in Earth's atmosphere, gaseous nitrogen is not readily available to most forms of life. Only a special subset of organisms are capable of turning gaseous nitrogen into forms usable for life. Some of the first organisms to do this were the cyanobacteria, which has led them down the path towards symbioses with various plants on many occasions. 

Cycads are but one branch of the gymnosperm tree. Their lineage arose at some point between the Carboniferous and Permian eras. Throughout their history it would seem that Cycads have done quite well in poor soils. They owe this success to a partnership they struck up with cyanobacteria. Although it is impossible to say when exactly this happened, all extant cycads we know of today maintain this symbiotic relationship with these tiny prokaryotic organisms. 

Cross section of a coralloid cycad root showing the green cyanobacteria inside. Photo by George Shepherd licensed under CC BY-NC-SA 2.0

The relationship takes place in Cycad roots. Cycads don't germinate with cyanobacteria in tow. They must acquire them from their immediate environment. To do so, they begin forming specialized structures called precoralloid roots. Unlike other roots that generally grow downwards, these roots grow upwards. They must situate themselves in the upper layer of soil where enough light penetrates for cyanobacteria to photosynthesize.

The cyanobacteria enter into the precoralloid roots through tiny cracks and take up residence. This causes a change in root development. The Cycad then initiates their development into true coralloid roots, which will house the cyanobacteria from that point on. Cycads appear to be in full control of the relationship, dolling out carbohydrates in return for nitrogen depending on the demands of their environment. Coralloid roots can shed and reform throughout the lifetime of the plant. It is quite remarkable to think about how nitrogen-fixing symbiotic relationships between plants and microbes have evolved independently throughout the history of life on this planet.

Photo Credits: [1] [2]

Further Reading: [1] [2]

The World's Only (Known) Photosynthetic Vertebrate

You may be asking yourself right now why I have posted a picture of a salamander this morning. This is a plant blog after all! Well, what I am about to tell you may seem a bit crazy, but I assure you this discovery has opened up some doors that science never really considered a possibility before. The yellow spotted salamander (Ambystoma maculatum) is the first and only (known) photosynthetic vertebrate ever discovered!

That's right. You heard me. A photosynthetic animal. More accurately speaking, it is the embryos of this species that undergo photosynthesis. To understand why this happens we must back up a little bit. Yellow spotted salamanders are a species of mole salamander that can be found in wet areas of eastern North America. They spend most of their adult lives underground, hiding beneath logs and rocks in the forest, feeding on any manner of invertebrates. Once a year (around this time) adult yellow spotted salamanders undertake a massive migration down to the pools where they mate. On the first few warm, rainy nights, thousands of salamanders can be seen trucking their way to vernal pools and ponds to breed. It is an amazing sight to behold.

The thing about yellow spotted salamanders is they will only breed in fishless ponds. Their larvae would be an easy meal for many predatory fish species. The problem that arises out of this breeding strategy is that fishless ponds tend to be very low in oxygen. It has long been known that the eggs of this species form a symbiotic relationship with an algae. The algae produce oxygen for the developing embryo and the embryo feeds the algae via its nitrogen rich waste and CO2. This relationship was always thought to be external, that is until Ryan Kerney of Dalhousie University in Halifax, Nova Scotia discovered that embryos of a certain age actually had algae living within their cells.

They algae don't seem to start off inside the cells though. This may be why this relationship wasn't discovered earlier. Roger Hangarter at Indiana University found that it isn't until parts of the salamander's nervous system begin to develop that the algae move into the embryo and set up shop. The algae then reside near the salamander's mitochondria, which are the powerhouses of the cell. So where are the algae coming from? While more research needs to be done, Karney also discovered the presence of algae in the oviducts of adult female spotted salamanders. It is looking like mother salamanders are actually passing the algae on to their offspring. 

Though this is the first and only instance we know of this sort of photosynthetic relationship in vertebrate animals, this discovery has opened the door for exploring the possibility of other photosynthetic symbionts. It has also allowed scientists a different avenue to explore just how cells recognize and deal with foreign bodies. We live in such an amazing world!

Further Reading: [1] [2] [3]

 

Orchid Dormancy Mediated by Fungi

Photo by NC Orchid licensed under CC BY-NC 2.0

North America's terrestrial orchids seem to have mastered the disappearing act. When stressed, these plants can enter into a vegetative dormancy, existing entirely underground for years until the right conditions return for them to grow and bloom. Cryptic dormancy periods like this can make assessing populations quite difficult. Orchids that were happy and flowering one year can be gone the next... and the next... and the next...

How and why this dormancy is triggered has confused ecologists and botanists alike. Certainly stress is a factor but what else triggers the plant into going dormant? According to a recent paper published in the American Journal of Botany, the answer is fungal.

Orchids are the poster children for mycorrhizal symbioses. Every aspect of an orchid's life is dependent on these fungal interactions. Despite our knowledge of the importance of mycorrhizal presence in orchid biology, no one had looked at how the abundance of mycorrhizal fungi influenced the life history of these charismatic plants until now.

By observing the presence and abundance of a family of orchid associated fungi known as Russulaceae, researchers found that the abundance of mycorrhizal fungi in the environment is directly related to whether or not an orchid will emerge. The team focused on a species of orchid known commonly as the small whorled pogonia (Isotria medeoloides). Populations of this federally threatened orchid are quite variable and assessing their numbers is difficult.

The team found that the abundance of mycorrhizal fungi is not only related to prior emergence of these plants but could also be used as a predictor of future emergence. This has major implications for orchid conservation overall. It's not enough to simply protect orchids, we must also protect the fungal communities they associate with.

Research like this highlights the need for a holistic habitat approach to conservation issues. So many species are partners in symbiotic relationships and we simply can't value one partner over the other. If conditions change to the point that they no longer favor the mycorrhizal partner, it stands to reason that it would only be a matter of years before the orchids disappeared for good.

Photo Credit: NC Orchid

Further Reading: [1]

Bacteria Help the Cobra Lily Subdue Prey

Photo by David Berry licensed under CC BY 2.0

The cobra lily (Darlingtonia californica) is one of North America's most stunning pitcher plants. Native to a small region between northern California and southwestern Oregon, this bizarrely beautiful carnivore lives out its life in nutrient poor, cold water bogs and seeps. Although it resides in the same family as our other North American pitcher plants, Sarraceniaceae, the cobra lily has a unique taxonomic position as the only member of its genus.

It doesn't take much familiarity with this plant to guess that it is carnivorous. Its highly modified leaves function as superb insect traps. Lured in by the brightly colored, tongue-like protrusions near the front tip of the hood, insects find a sweet surprise. These tongue-like structures secrete nectar. As insects gradually make their way up the tongue, they inevitably find themselves within the downward pointing mouth of the pitcher. This is where those translucent spots on the top of the hood come in.

Those translucent spots trick the insects into flying upwards into the light. Instead of a clean getaway, insects crash into the inside of the hood and fall down within the trap. The slippery walls of the pitcher interior make escape nearly impossible but that isn't the only thing keeping insects inside. Research has shown that the cobra lily gets a helping hand from bacteria living within the pitcher fluid.

Unlike other pitcher plants, the cobra lily does not fill its traps with rain water. The downward pointing mouth prevents that from happening. Instead, the pitchers secrete their own fluid by pumping water up from the roots. Although there is evidence that the cobra lily does produce at least some of its own digestive enzymes, it is largely believed that this species relies heavily on a robust microbial community living within its pitchers to do most of the digesting for it. This mutualistic community of microbes saves the plant a lot of energy while also providing it with essential nutrients like nitrogen in return for a safe place to live.

That isn't all the bacteria are doing for this pitcher plant either. As it turns out, the pitchers' microbial community may also be helping the plant capture and subdue its prey. A study based out of UC Berkeley demonstrated that the presence of these microbes helps lower the surface tension of the water, effectively drowning any insect almost immediately.

Some members of the microbial community release special compounds called biosurfactants. Through an interesting chemical/physical process that I won't go into here, this keeps insects from using the surface tension of the water to keep them afloat, not unlike a water strider on a pond. Instead, as soon as insects hit the bacteria infested waters, they break the surface tension and sink down to the bottom of the pitcher where they quickly drown. There is little chance of escape for a hapless insect unlucky enough to fall into a cobra lily trap.

Although plant-microbe interactions are nothing new to science, this example is the first of its kind. Although this prey capture role is very likely a secondary benefit of the microbial community within the pitchers, it certainly makes a big difference for these carnivores living in such nutrient poor conditions.

Read more about the amazing world of carnivorous plants by picking up a copy of my book!

Photo Credit: [1] [2]

Further Reading: [1]

Live-In Mites

Photo by Scott Zona licensed under CC BY-NC 2.0

Hearing the word "mite" as a gardener instantly makes me think of pests such as spider mites. This is not fair. The family to which mites belong (Acari) is highly varied and contains many beneficial species. Many mites are important predators at the micro scale. Some are fungivorous, eating potentially harmful species of fungi. Whereas this may be lost on the majority of us humans, it is certainly not lost on many species of plants. In fact, the relationship between some plants and mites is so strong that these plants go as far as to provide them with a sort of home.

Photo by Jsarratt licensed under CC BY-SA 3.0

Domatia are specialized structures that are produced by plants to house arthropods. A lot of different plant species produce domatia but not all of them are readily apparent to us. For instance, many trees and vines such as red oak (Quercus rubra), sugar maples (Acer saccharum), black cherries (Prunus serotina), and many species of grape (Vitis spp.) produce tiny domatia specifically for mites. The domatia are often small, hairy, and function as shelter for both the mites and their eggs.

By housing certain species of mites, these plants are ensuring that they have a steady supply of hunters and cleaners living on their leaves. Predatory mites are voracious hunters, keeping valuable leaves free of microscopic herbivores while frugivorous mites clean the leaves of detrimental fungi that are known to cause infections such as powdery mildew. The exchange is pretty straight forward. Mites get a home and a place to breed and the plants get some protection. Still, some plants seem to want to sweeten the relationship in a literal sense.

Some plants, specifically grape vines in the genus Vitis, also produce extrafloral nectaries on their leaves. These tiny glands secrete sugary nectar. In a paper recently published in the Annals of Botany, it was found that extrafloral nectaries enhances the efficacy of these mite domatia by enticing more mites to stick around. By adding nectar to domatia-producing leaves that did not secrete it, the researchers found that nectar increases beneficial mite densities on these leaves by 60 - 80%. This translates to an increase in fitness for these plants in the long run.

I love research like this. I had no idea that so many of my favorite and most familiar tree and vine species had entered into an evolutionary relationship with beneficial mites. This adds a whole new layer of complexity to the interactions within any given environment. It just goes to show you how much is left to be discovered in our own back yards.

Photo Credits: [1] [2] [3]

Further Reading: [1] [2] [3]

The Whorled Pogonia

I live for moments like this. The only downside to that is I can never really predict when they are going to happen. There I was driving up a mountain road in search of a handful of other plant species related to my research. The road was narrow and there was a steep bank on the drivers side. The Southern Appalachian Mountains are brimming with botanical diversity. As such, it can be hard to tease out individual plants, especially while driving. This is why having a refined search image comes in handy. 

I was rounding a bend in the road when something out my window caught my eye. My mind went racing and it wasn't long before a suspicion crept into my head. If I was right, this was an opportunity I was not going to miss. I found the nearest pull off, parked the truck, and ran back down the road. I am so happy that I decided to trust my instincts. There in front of me was a small population of whorled pogonia orchids (Isotria verticillata). 

It was like being in the presence of a celebrity that I had been stalking for years. This was an orchid I have been dying to see. The harder I looked the more I saw. I had to sit down. Here in front of me was a species of orchid that isn't seen by many. In fact, entire populations of these species can go unseen for decades until they have enough energy to flower. 

Flowering in this species is said to be quite erratic. Because they live in shaded environments, building up the energy needed to reproduce can be difficult. Like all orchids, the whorled pogonia relies on an obligate relationship with mycorrhizal fungi to supply the nutrients it needs. In return, the orchids provide fungi with carbohydrates. The problem with erratic flowering, however, is that it makes reproduction difficult. Rarely are two populations flowering at the same time and in close enough proximity for successful cross pollination. More often, these orchids will self fertilize, which can lead to high rates of inbreeding. 

Large bees are the main pollinators of the whorled pogonia. The flowers themselves are reported to produce a feint odor reminiscent of Vanilla. This is interesting to note because in the greater scheme of orchid phylogenetics, this species is placed in the Vanilla subfamily, although such distinctions can get muddled quickly. Regardless, simply being in the presence of this orchid was enough to give me goosebumps. It is a shame that such a species is being lost throughout much of its range. 

Further Reading:
http://bit.ly/1ssBmdF

http://bit.ly/1WEmZzm

On Peonies and Ants

Photo by George Vopal licensed under CC BY 2.0

It is just about that time when peony buds burst forth and put on their late spring display. My mother loves her peonies and she gets very excited every year when they bloom. It's adorable. However, she has always been disgusted by the amount of ants the peonies attract. Indeed, many people all over the internet seem to feel the same way. Growing up, I always wondered why the ants seemed to swarm all over peony buds, so I decided to look into it a little deeper.

There are many sources out there that claim that peonies need ants in order to bloom. To me, this seems very maladaptive on the part of the peony. The genus Paeonia is represented in Asia, Southern Europe and parts of western North America. I am going to assume that the ant/peony relationship didn't start in the garden so it's roots have to be somewhere in the evolutionary history of the plant. What sense does it make for a plant to produce flower buds that excrete sticky sugars that keep them from opening until something cleans the sugars off? In fact, despite anecdotal reports, peony buds will open without ants. So then why does the plant bother to produce sugars that attract ants?

Interestingly enough, despite a good amount of searching, there is not a lot of research done on this subject but the answer to this question can come from looking at how ants interact with other plants and animals. Many plant species have special glands on their stems that produce sugary secretions which attract ants. It's not just plants either. Insects such as aphids and leafhoppers famously excrete honeydew that ants can't resist. In each of these cases, organisms are using the ants' natural tendency to guard a food source. The ants will viciously attack anything that threatens this easy meal.

It would seem to me that the peonies are doing just that with their flower buds. By secreting a sugary substance during their development, the plant are likely recruiting ants to protect the flowers, which for angiosperms, are the most precious part of the plant. It takes a lot out of a plant to flower and the threat of herbivory is ever present. If an insect tries to take a bite out of a bud, the ants quickly swarm and drive it off. It's a win-win situation. The ants get an easy, high-energy food source and the plant suffers less damage to its reproductive organs.

The scary part to me in researching all of this is plethora of information out there on how to get rid of the ants. People go through chemical after chemical to rid their peonies of ants when, in reality, the ants are some of the best friends a peony could have! So leave those ants alone and enjoy the free pest removal services they provide every spring!

Photo Credit: [1]

Further Reading:

http://www.youtube.com/watch?v=Gnm2nV_nwOk

Of Gunnera and Cyanobacteria

Photo by UnconventionalEmma licensed under CC BY-NC 2.0

Nitrogen is a limiting resource for plants. It is essential for life functions and yet they do not produce it on their own. Instead, plants need to get it from their environment. They cannot uptake gaseous nitrogen, which is a shame because it makes up 78.09% of our atmosphere. As such, some plants have developed very interesting ways of obtaining nitrogen from their environment. Some, like the legumes, produce special nodules on their roots, which house bacteria that fix atmospheric nitrogen. Other plants utilize certain species of mycorrhizal fungi. One family of plants, however, has evolved a symbiotic relationship that is unlike any other in the angiosperm world.

A Gunnera inflorescence. Photo by Lotus Johnson licensed under CC BY-NC 2.0

Meet the Gunneras. This genus has a family all to itself - Gunneraceae. They can be found in many tropical regions from South America to Africa and New Zealand. Some species of Gunnera are small while others, like Gunnera manicata, have leaves that can be upwards of 6 feet in diameter. Their leaves are well armed with spikes and spines. All in all they are rather prehistoric looking. The real interesting thing about the Gunneras though, is in the symbiotic relationship they have formed with cyanobacteria in the genus Nostoc.

Traverse section of a Gunnera stem showing cyanobacteria colonies (C) and the cup-like structures (S) where they enter the stem. [SOURCE]

Gunnera produce cuo-like glands that house these cyanobacteria. The glands are filled with a special mucilage that not only attracts the cyanobacteria, but also stimulates it to grow. Once inside the glands, the cyanobacteria begins to grow into the plant, eventually fusing with the Gunnera cells. From there the cyanobacteria earn their keep by producing copious amounts of usable nitrogen and in return, the Gunnera supplies carbohydrates. This relationship is amazing and quite complex. It also offers researchers an insight into how such symbiotic relationships evolve.

Photo Credit: [1] [2] [3]

Further Reading: [1] [2] [3] [4] [5]

Sticky Friend

Photo by David A. Hofmann licensed under CC BY-NC-ND 2.0

We have all had encounters with sticky plants. Outside of being an interesting sensory experience, the sticky nature of these floral entities would appear to have some evolutionary significance. Considering the cost of producing the glandular trichomes responsible for their stickiness, function is a reasonable question to ask about. For anyone who has taken the time to observe such plants, you will have undoubtedly noticed that insects tend to get stuck to them.

For carnivorous plants, the utility of these glands is readily obvious - trapped insects become food. Even non-carnivores like Roridula gain a nutrient benefit in the form of nutrient-rich feces deposited around the plant by specialized carnivorous bugs that consume trapped insects. However, there are many species of plants out there that fall under the category of "sticky" and a new paper explores this in a more general way.

The serpentine columbine (Aquilegia eximia) is endemic to the Coastal Range of California and it is indeed quite sticky. Its surfaces are covered in glandular hairs. Any given plant can be covered in insects unfortunate enough to come into contact with it. However, it is not a carnivore. As such, researchers wanted to see what benefits, if any, the columbine gained from producing these glands.

By manipulating the amount of insects that were stuck to each plant, researchers found that plants without "victims" actually received more insect damage. The key to this mystery were predators. Plants with lots of trapped victims had more predatory bugs hanging around. These predators, when present, reduced herbivory by deterring other insects that were too large to get stuck. What's more, most of the benefits were observed in the flower buds, which means predators increased the overall reproductive fitness of the serpentine columbine. If the columbine did not trap small insects, these predators would have no reason to hang around.

These predatory bugs were by no means specific to the columbine. In fact, observation of the surrounding plant community found that these predatory insects were present on other sticky genera such as Arctostaphylos, Hemizoni, Holocarpha, Calycidenia, Cordelanthus, Castilleja, Mimulus, Trichostema, and Grindelia. This suggests that the relationship between sticky plants and these generalist predators is more widespread than previously thought. It may also offer a unique window into one possible driver behind the evolution of carnivory in plants.

Photo Credit: David A. Hofmann (http://bit.ly/1l9OtwC)

Further Reading:
http://www.esajournals.org/doi/abs/10.1890/15-0342.1

Echoes of a Glacial Past

Climate change is often talked about in the context of direct effects on species. However, as John Muir so eloquently put it, "When we try to pick out anything by itself, we find it hitched to everything else in the Universe." In essence, nothing is ever black and white and the research I am writing about today illustrates this fact quite well.

Ants and plants have some very intricate interactions. A multitude of plant species rely on ants as their seed dispersers. Many of these plant species are spring ephemerals that take advantage of the fact that there is little else for ants to eat in the early spring by attaching fatty capsules to their seeds that are very attractive to foraging ant species. We refer to seed dispersal by ants as “myrmecochory.”

There are two big players in the foraging ant communities of eastern North America, the warm adapted Aphaenogaster rudis and the cold adapted Aphaenogaster picea. The cold adapted A. picea emerges from winter dormancy early in the spring while the warm adapted species emerges from dormancy much later in the spring. In the southern portions of their range, A. rudis outcompetes A. picea.

What is the big deal? Well, the researchers looked at two plant species that rely on these ants for seed dispersal, Hepatica nobilis and Hexastylis arifolia. Hepatica nobilis sets seed early in the spring, relying on ant species like A. picea to disperse its seed whereas Hexastylis arifolia sets seed late in spring, which is prime time for A. rudis. Researchers noticed that, in the southern portions of their range where A. picea had been displaced, Hepatica has a very clumped and patchy growth habit where farther north it did not. Hexastylis on the other hand seemed to have a more normal growth pattern in the south.

By performing some transplanting experiments and examining foraging and seed dispersal, they found that the absence of A. picea in the south spelled ecological disaster for Hepatica. It continues to set seed but because A. rudis emerges long after seed set, it is not filling the gap left by the missing A. picea. Hexastylis, which only grows in the south and sets seed much later, does just fine with the warm adapted A. rudis. Farther north where A. picea still rules, Hepatica has no trouble with seed dispersal but Hexastylis drops out of the ecosystem entirely. In essence, because of warming climate trends since the end of the Pleistocene, Hepatica is falling out of sync with its mutualistic ant partner in the southern portions of its range and, in time, may become extirpated.

Further Reading: [1]