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]

Meet the Sweetfern

Photo by Sten Porse licensed under CC BY-SA 3.0

Photo by Sten Porse licensed under CC BY-SA 3.0

I remember the first time I laid my eyes on Comptonia peregrina. I was new to botany at that point in my life so I didn't have a well developed search image for these sorts of things. I was scrambling down a dry ridge with a scattered overstory of gnarly looking chestnut oaks when I saw a streak of green just below me on a sandy outcropping. They were odd looking plants, the likes of which I had never seen before.

I took out my binoculars to get a better look. What were these strange organisms? Were they ferns? No, they seemed to have woody stems. Were they gymnosperms? No, I could make out what appeared to be male catkins. Luckily I never leave home without a field guide or two. Using what little terminology I knew, I was able to narrow my focus to a plant commonly called a "sweetfern."

Photo by Megan Hansen licensed under CC BY-SA 2.0

Photo by Megan Hansen licensed under CC BY-SA 2.0

This was one of the first instances in which I grasped just how troublesome common names can be. C. peregrina is mostly definitely not a fern. It is actually an angiosperm that hails from the bay family (Myricaceae). Comptonia is a monotypic genus, with C. peregrina being the only species. It is a denizen of dry, nutrient poor habitats. As such, it has some wonderful adaptations to deal with these conditions.

To start with, its a nitrogen fixer. Similar to legumes, it forms nodules on its roots that house specialized nitrogen-fixing bacteria called rhizobia. This partnership takes care of its nitrogen needs, but what about others? One study found that not only do the roots form nodules, they also form dense cluster roots. Oddly, closer observation found that these clusters were not associated with mycorrhizal fungi. What's more, they also found that these structures were most prevalent in highly disturbed soils. It is thought that this is one way that the plant can maximize its uptake of phosphorus under the harshest growing conditions. 

Photo by Jomegat licensed under CC BY-SA 3.0

Flowering in this species is not a showy event. C. peregrina can be monoecious or dioecious, producing male and female catkins towards the ends of its shoots. After fertilization, seeds develop inside bristly fruits. Seed banking appears to be an important reproductive strategy for this species. One study found that germinated seeds had lain dormant in the soil for over 70 years until disturbance opened up the canopy above. It is expected that seeds of this species could exhibit dormancy periods of a century or more. 

In total, this is one spectacular species. Not only does it have a unique appearance, it is also extremely hardy and an excellent species to plant in drought-prone soils wherever it is native. I do see it in landscaping from time to time. If you encounter this species in the wild, take the time to observe it in detail. You will be happy you did!

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

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

Going Veg With Nepenthes ampullaria

Photo by Bernard DUPONT licensed under CC BY-SA 2.0

Photo by Bernard DUPONT licensed under CC BY-SA 2.0

Carnivory in the plant kingdom is an interesting evolutionary adaptation to living in nutrient poor environments. It has arisen in only a handful of different plant families and indeed, the genera that exhibit it are considered highly derived. There is something to be said about a sessile organism that can take down mobile prey at the rate that most carnivorous plants do.

Perhaps part of our fascination with these botanical wonders stems from their move towards dietary habits not unlike our own. The reason for their predatory behavior is to acquire nutrients like nitrogen and phosphorus. Without these essential nutrients, life as we know it would not exist. It is no wonder then that carnivorous plants have evolved some very interesting ways of getting them into their tissues and to me, there is nothing more peculiar than the way in which Nepenthes ampullaria gets its much needed nitrogen fix.

A rather widespread species, N. ampullaria is at home in the understory of the rain forests of the southeast Asian islands. It differs from its carnivorous cousins in a multitude of ways. For starters, the pitchers of N. ampullaria are oddly shaped. Resembling an urn, they sit in dense clusters all over the jungle floor, below the rest of the plant. Unlike other Nepenthes, the pitchers have only a small, vestigial lid with no nectar glands. Finally, the slippery, waxy surface that normally coats the inside of most Nepenthes pitchers is absent in the pitchers of N. ampullaria. All of these traits are clues to the unique way in which this species has evolved to acquire nitrogen.

N. ampullaria doesn't lure and digest insects. Instead, it relies on leaf litter from the forest canopy above for its nutritional needs. The urn-like shape, lack of a hood, and clustered growth enable the pitchers to accumulate considerable amounts of leaf litter in the pitchers. Because the pitchers are relatively long lived for a Nepenthes, lasting upwards of 6 months, they offer up a nice microhabitat for a multitude of insect and even frog larvae. The collective group of organisms living within the pitchers are referred to as an inquiline community.

Over time, an inquiline community develops in each of the pitchers. This is the key to the success of N. ampullaria. As the inquiline organisms breakdown the leaf litter, they release copious amounts of nitrogen-rich waste. The pitchers can then absorb this waste and begin to utilize it. At least one study found that an individual plant can obtain 35.7% of its foliar nitrogen in this manner. It has also been demonstrated that the pitchers actively manipulate the pumping of hydrogen ions into the fluid within to keep it less acidic than that of other Nepenthes.

I don't know if I would consider this a case of herbivory as the nitrogen is still coming from an animal source but it is nonetheless an interesting adaptation. Instead of using valuable resources on actively digesting its own prey, N. ampullaria is getting other organisms to do the work for it. Not too shabby.

Further Reading:

http://bit.ly/1IRbYG9

http://jxb.oxfordjournals.org/content/61/5/1365

http://link.springer.com/article/10.1007/s004420050390

http://bit.ly/1S10oej