Insect Killer, Plant Symbiont

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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)

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

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

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

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

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.

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]

Mutant Orchids Have a lot to Teach Us About Parasitic Plants

A) Albino and (B) green individual of Goodyera velutina.

A) Albino and (B) green individual of Goodyera velutina.

The botanical world is synonymous with the idea of photosynthesis. Plants take in carbon dioxide and water and utilize light to make their own food. However, not all plants make a living this way. There are many different species of plants that have evolved a parasitic lifestyle to one degree or another. Some of my favorites are those that parasitize mycorrhizal fungi. We call these plants “mycoheterotrophs” and they are fascinating to say the least. Orchids are especially prone to this strategy, with over 1% of all known species having completely lost the ability to photosynthesize.

Our knowledge of the mycoheterotrophic strategy is fragmentary at best. We still don’t fully understand things like how the plants obtain what they need from the fungus nor how they are able to maintain their parasitic lifestyle without the fungus catching on and rejecting the one-sided partnership. This is not to say we know nothing. In fact, as technologies advance, we are unlocking at least some of the mysteries of mycoheterotrophic plants. Some of the best advances come from studying mutant, albino orchids. To understand how, we have to take a closer look at the “average” orchid lifestyle.

Orchids in general make great candidates for understanding the evolution of mycoheterotrophy because all of them start their lives as parasites. Orchids produce some of the smallest seeds in the plant kingdom and without the help of mycorrhizal fungi, they would never be able to germinate. For much of their early life, orchids rely on fungi to provide them with both their mineral and carbohydrate needs. Only after the orchids are large enough to grow leaves will most of them start to give back to their fungal partners in the form of carbohydrates generated from photosynthesis.

Still, many orchids never fully let go of this parasitic lifestyle. This is especially true for orchids living under dense forest canopies. With light in limited supply, many orchids adopt a mixotrophic lifestyle. Essentially this means that although they actively photosynthesize, they nonetheless rely on fungi to provide them with both carbohydrates and minerals. Mixotrphy is likely the most wide-spread orchid strategy and it has been hypothesized that it is also the first step along the path to becoming fully parasitic. This is where the mutant orchids enter the equation.

(A) Albino and (B) green individuals of Epipactis helleborine

(A) Albino and (B) green individuals of Epipactis helleborine

Every once in a while, some orchids will germinate and grow into albino versions of their species. Without the ability to produce chlorophyll, these mutants should be destined for a quick death. Such is not the case for many of these orchids. Albino orchids often go on to live full lives, growing and flowering just like their photosynthetic progenitors. Although they do exhibit signs of reduced fitness, the fact that they are able to live at all brings up a lot of questions ready for science to tackle.

Recent investigations into the lives of these albino mutants has revealed some interesting insights into how mycoheterotrophy may have evolved in the first place. By studying the fungal partners of both healthy plants and the albinos, researchers have been able to demonstrate that albinos are doing things a bit differently than their photosynthetic parents. Using isotopes of carbon and nitrogen, scientists are discovering that the albinos have switched their interaction with the fungi in such a way that they more resemble fully mycoheterotrophic orchid species. This is done despite the fact that both albinos and their fully functional parents associate with the same guild of mycorrhizal fungi.

Another interesting difference between albinos and their photosynthetic parents is the fact that the genes involved both antioxidant metabolism and metabolite transfer (mainly carbon in this case) were more active in the albinos than they were in functioning plants. The uptick in gene functioning related to antioxidant metabolism suggests that the mutant plants are undergoing greater oxidative stress than their functional parents. This may have something to do with how the albinos are able to obtain nutrients from their fungal partners. It is thought that mycoheterotrophs actively digest parts of the fungi, which allows them to access the carbon and minerals they need to survive. This process exposes their cells to reactive oxygen compounds that can be very damaging. Antioxidants would help to reduce such damage.

The uptick in genes associated with metabolite transfer was more surprising because it suggests that despite being parasites, the plants are actively transferring substances back to the fungi. It has long been assumed that mycoheterotrophy was a one way street, with fungi transferring nutrients to plants only. These genes now suggest that, at least in some species, such transfer is a two-way street. The exact nature of this two-way transfer remains a mystery and certainly brings up many more questions that must be asked before we can better understand this relationship.

All of this is not to say that such albino mutants are fruitful “next steps” in the evolution of these species. Far from it, in fact. Two things that most albino orchid variants have in common is the fact that they are rare and, of those that have been studied, produce far fewer seeds. There are a lot of anatomical and physiological differences between true mycoheterotrophic species and albino variants and it appears that without those anatomical adaptations, the albinos are a lot less fit than their photosynthetic parents. As authors Selosse and Roy put it:

“non-chlorophyllous variants are likely to represent unique snapshots of failed transitions from mixotrophy to mycoheterotrophy. They are ecological equivalents to mutants in genetics, that is, their dysfunctions might suggest what makes mycoheterotrophy successful. Although their determinism remains unknown, they offer fascinating models for comparing the physiology of mixo- and mycoheterotrophs within similar genetic backgrounds.”

Mutants are strange indeed but with the right kinds of questions and approaches, they have a lot to teach us about ecology, evolution, and life at large.

Photo Credits: [1] [2]

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