The Bladderwort Microbiome Revealed

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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.

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. 

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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] [4]

Further Reading: [1]

 

The Tiniest Flowers

It is easy to get caught up in grandeur. Finding the largest of something is always fun but what about the smallest? I have talked about the largest flowers in the world but how small can a flower get? The answer lies in some of the most common plants on earth, the duckweeds (Wolffia spp. & Lemna spp.).

Duckweeds represent the epitome of flowering plant reductionism. They are little more than a pair or so of leaf-like structures that float at or near the water’s surface and the occasional taproot. What is even more striking are their flowers. Whereas most duckweed will reproduce clonally, every once in a while a flower will be produced. One species, Wolffia globosa, produces the smallest flowers of any duckweed. They are also the smallest flowers known to science. As you can see from the picture, they are nothing more than a couple anthers (A1 and A2) and a pistil (Pi). What's more, the male parts mature and die before the female parts, thus limiting the chances of self fertilization. As one would expect, the fruits produced are also the smallest fruits in the world.

Wolffia globosa

Wolffia globosa

This is all quite interesting considering the fact that DNA analysis places duckweeds in the aroid family (Araceae), which is home to the largest single inflorescence in the world! That's right, the arum family produces one of the largest groups of flowers in the world as well as the smallest flower and fruit in the world. Another cool fact about duckweed, specifically Wolffia, is that, gram for gram, it produces the same amount of protein as soy beans. Because of this, it is used as a food plant in many places around the world and more countries are waking up to its potential as both a food plant and a phytoremediation species. Pretty neat for such a small plant.

Photo Credit: Patrick Denny

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

Aquatic Angiosperm: A Cretaceous Origin?

It would seem that yet another piece of the evolutionary puzzle that are flowering plants has been found. I have discussed the paleontological debate centered around the angiosperm lineage in the past (http://bit.ly/1S6WLkf), and I don't think the recent news will put any of it to rest. However, I do think it serves to expand our limited view into the history of flowering plant evolution.

Meet Montsechia vidalii, an extinct species that offers tantalizing evidence that flowering plants were kicking around some 130–125 million years ago, during the early days of the Cretaceous. It is by no means showy and I myself would have a hard time distinguishing its reproductive structures as flowers yet that is indeed what they are thought to be. Detailed (and I mean detailed) analyses of over 1,000 fossilized specimens reveals that the seeds are enclosed in tissue, a true hallmark of the angiosperm lineage.

On top of this feature, the fossils also offer clues to the kind of habitat Montsechia would have been found in. As it turns out, this was an aquatic species. The flowers, instead of poking above the water, would have remained submerged. An opening at the top of each flower would have allowed pollen to float inside for fertilization. Another interesting feature of Montsechia is that it had no roots. Instead, it likely floated around in shallow water.

This is all very similar to another group of extant aquatic flowering plants in the genus Ceratophyllum (often called hornworts or coon's tail). Based on such morphological evidence, it has been agreed that these two groups represent early stem lineages of the angiosperm tree. Coupled with what we now know about the habitat of Archaefructus (http://bit.ly/1S6WLkf), it is becoming evident that the evolution of flowers may have happened in and around water. This in turn brings up many more questions regarding the selective pressures that led to flowers.

What is even more amazing is that these fossils are by no means recent discoveries. They were part of a collection that was excavated in Spain over 100 years ago. Discoveries like this happen all the time. Someone finds a interesting set of fossils that are then stored away on a dark shelf in the bowels of a museum only to be rediscovered decades or even centuries later.

All in all I think this discovery lends credence to the idea that flowering plants are a bit older than we like to think. Also, one should be wary of anyone claiming to have found "the first flower." The idea that there could be a fossil out there that depicts the first anything is flawed a leads to a lot of confusion. Instead, fossils like these represent snapshots in the continuum that is evolution. Each new discovery reveals a little bit more about the evolution of that lineage. We will never find the first flower but we will continue to refine our understanding of life on this planet.

Photo Credits: Bernard Gomeza, Véronique Daviero-Gomeza, Clément Coiffardb, Carles Martín-Closasc, David L. Dilcherd, and O. Sanisidro,

Further Reading:
http://www.pnas.org/content/112/35/10985.abstract

Aquarium Banana

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One of my first true passions in life was maintaining freshwater aquariums. There is something about being able to observe a world totally foreign to our own that drew me in. It wasn't long before I discovered the splendor of planted aquascapes. I would have to say that my first foray into this realm probably planted the botanical seed that would later explode into the obsession it is today. 

I was always rather perplexed by a plant that I would see for sale at the local aquarium shop. They were labelled as "banana plants" because of their peculiar root structures. They never seemed to fit my aesthetic in those early days so I largely passed them by. Recently I have gotten back into aquariums, only this time it is very plant centric. While perusing the plants offered here in town, I again came across the peculiar banana plant. 

This time around, I am a bit more versed in taxonomy and this plant made more sense. I realized that the banana plants we see for sale for aquariums are small, immature specimens of some sort of "lily pad." A deeper investigation would prove me correct. Though not a true water lily (family Nymphaeaceae), the banana plants nonetheless take on a similar growth form with large, floating leaves emerging from an underwater rootstock. 

Banana plants are known scientifically as Nymphoides aquatica. Their generic name comes from their striking similarity to the afore mentioned water lilies. However, this resemblance is merely superficial. Banana plants belong to the family Menyanthaceae, making it a relative of plants like buckbean (Menyanthes trifoliata). They grow native in calm bodies of water throughout southeastern North America. Whereas the young leaves grow immersed, larger adult leaves eventually make their way to the surface where they float. 

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From time to time, small white flowers are produced. This is when its familiar affiliation makes the most sense. This species is dioecious, though seed set is apparently sporadic. Regardless, banana plants readily reproduce vegetatively, either by fragmentation of their roots or by broken leaves settling in a spot and forming roots themselves. 

So far this is an interesting aquarium specimen. It seems to have adjusted to my aquarium rather well and it grows pretty quickly. In time I hope it performs more like it does in the wild than as a sad, stunted specimen doomed to a slow death. Only time will tell. 

Flower Pic: Show_ryu (Wikimedia Commons)

Further Reading: [1]