Should We Be Calling Aquatic Bladderworts Omnivores Instead of Carnivores?

Photo by Leonhard Lenz licensed under CC BY-NC 2.0

As is so often the case in nature, the closer we start to look at things, the more interesting they become. Take, for instance, the diet of some carnivorous bladderworts (Utricularia spp.). These wonderful organisms cover their photosynthetic tissues in tiny bladder traps that rapidly spring open whenever a hapless invertebrate makes the mistake of coming too close to a trigger hair. The unlucky prey is quickly sucked into the trap and subsequently digested.

This is how most bladderworts supplement their growth. As cool as this mechanism truly is, our obsession with the idea that these plants are strict carnivores has historically biased the kinds of investigations scientists attempt with these plants. Over the last decade or so, closer inspection of the contents of aquatic bladderwort traps has revealed that a surprising amount of plant material gets trapped as well. Most of this material consists of single celled algae. Is it possible that at least some aquatic bladderworts also gain nutrients from all of that “vegetable” matter?

The answer to this question is a bit more nuanced than expected. Yes, it does appear that non-animal material frequently ends up in bladderwort traps. Much of this comes in the form of a wide variety of algae species. What’s more, it does appear that algae are broken down within the traps themselves, suggesting that the bladderworts are actively digesting this material. The main question that needs to be answered here is whether or not the bladderworts actually benefit from the breakdown of algae.

Evidence of a nutritive benefit from algae digestion is mixed. Some studies have found that the bladderworts don’t appear to benefit at all from the breakdown of algae within their traps. Alternatively, others have found that bladderworts may benefit from digesting at least some types of algae. These authors noted that there doesn’t seem to be any benefit in terms of additional nitrogen for the bladderwort but instead suggest that other trace nutrients might be obtained in this way.

One of the biggest hurdles in this line of research arises from the fact that we still don’t fully understand the digestive mechanisms of bladderworts. It is possible that some of the algal degradation within bladderwort traps has nothing to do with digestion at all. Instead, it could simply be that algae stuck in the traps eventually dies and rots away. Another major question raised by these observations is how tiny organisms like single celled algae even make it into the traps in the first place. What we can say for sure is most algae are far too small to actually trigger the bladder traps. As such, algae is either getting into the traps passively via some form of diffusion or they are sucked in when other, larger prey is captured.

Some research has even suggested that the benefit of trapping algae may depend on the habitats in which bladderworts are growing. Bladderworts living in more acidic water have shown to capture far more algae than bladderworts in more neutral or alkaline water. This has to do with acidity. Numerically speaking, there is far less zooplankton living in acidic water than algae, which means algae is more likely to end up in the bladders. It could be that the benefits of algae are thus greater for plants living in places where little zooplankton is available. Certainly more work will be needed before we can call bladderworts omnivores but the idea itself is exciting.

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

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

Of Bladderworts & Birds

Photo by Jean and Fred licensed by CC BY 2.0

Bladderworts are as beautiful as they are deadly. Though they are known the world over for their carnivorous bladder traps, their flowers are something to marvel at as well. Bladderworts flower in a range of colors from yellows to whites, purples to reds. What’s more, the variety of shapes and sizes among bladderwort flowers are incredible. Though the vast majority of bladderwort species rely on insects for pollination, at least one species appears to have co-opted a bird for its reproductive needs.

Red coats (Utricularia menziesii) are endemic to a few coastal regions of western Australia. They are not floating aquatic plants like many of their North American cousins, nor do they grow epiphytically like many tropical bladderworts. Red coats are terrestrial in their habit. Moreover, they live in habitats that dry up for good portions of the year. As the soils dry out, red coats die back into tiny corms in which they store energy during their dry dormancy that will fuel growth as soon as rains return and the surrounding soils are once again saturated.

Photo by Jean and Fred licensed by CC BY 2.0

When conditions are right, red coats produce some of the most spectacular flowers of the entire genus. Though other species also produce red flowers, few produce such outlandishly bright blossoms. Moreover, the flowers themselves are rather robust structures complete with a long, tough nectar spur. Their color, form, and proximity to the ground has led more than one author to suggest that birds, not insects, are the main pollinators of this species.

Indeed, it appears that birds are what these flowers are attracting. Not just any bird will do either. It seems that the western spinebill (Acanthorhynchus superciliosus) is wonderfully primed to pollinate this lovely little carnivore. Red is a major attractant for birds and the fact that red coat flowers are presented so close to ground level places at the perfect height for ground-foraging spinebills. Also, the length, curvature, and nectar content of the nectar spur fits the spinebill beak nicely. Birds approach the plants on the ground and dip their long, curved beaks into the flower, picking up and depositing pollen as they go.

The western spinebill (Acanthorhynchus superciliosus). Note the curved beak. Photo by Jean and Fred licensed by CC BY 2.0

This isn’t the only bladderwort to be suspected of bird pollination. At least two others (Utricularia quelchii & Utricularia campbelliana) have been hypothesized to utilize hummingbirds for pollination. However, there is scant evidence for this. Pollination studies can be tricky like that. Without proper observation and study, one simply can’t confirm a particular pollination syndrome.

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

Further Reading: [1]

The Floating Bladderwort

A carnivorous plant species that uses its radially arranged stolons like tiny pontoons to float at the waters surface may sound like something out of a science fiction novel. However, it is a very real strategy  adopted by one of the coolest carnivorous plants in North America. Utricularia inflata is one of the largest species of floating bladderwort on this continent and it is a species worth knowing.

Sometimes referred to as the swollen bladderwort, this species enjoys a native range that extends through much of the southeastern United States. For most of the year it exists in a state quite similar to other aquatic bladderworts. It has no true roots or leaves. Instead it produces a long, filiform stolon covered in tiny filaments that act as leaves with bladder traps situated at their tips. It sits in the water  column, gobbling up anything small and unfortunate enough to stumble into it.

Photo by Daiju Azuma licensed under CC BY-SA 2.5

When flowering time approaches, these aquatic carnivores begin producing a different kind of stolon. Arranged like spokes on a wheel, the plant puts out swollen, air-filled stolons that float at the waters surface. These structures support the inflorescence. Flowers are bright yellow and resemble those of many other bladderwort species. Entire bodies of water can literally erupt in a sea of yellow bladderwort flowers when the right conditions present themselves.

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Photo by Adam Arendell licensed under CC BY-NC 2.0

As mentioned, this species is carnivorous. It uses tiny bladder traps to suck in unsuspecting prey. Their diet is varied and includes pretty much anything that can fit into its bladder traps. One research paper reports both animal (rotifers, cladocerans, copepods, annelids, rhizopodeans, as well as small insects) and "plant" (Bacillariophyta, Chlorophyta, Cyanophyta, and Euglenophyta) prey.

Unfortunately these plants have been introduced far outside of their native range. In many areas they are becoming prevalent enough to be considered invasive. For instance, research done in the Adirondack Mountains of New York found that the presence of introduced populations of U. inflata caused significant changes in nutrient cycling, sediment chemistry, and overall net primary productivity.

This is a very neat species well worth a closer look. That being said, if you are a hobbyist such as myself, it is very important to remember that we should never release a species (no matter how cool it is) into areas where it isn't native.

Photo Credit: Dr. Mark Whitten, [3] [4]

Further Reading: [1] [2]

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]