How Aroids Turn Up the Heat

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A subset of plants have evolved the ability to produce heat, a fact that may come as a surprise to many reading this. The undisputed champions of botanical thermogenesis are the aroids (Araceae). Exactly why they do so is still the subject of scientific debate but the means by which heat is produced is absolutely fascinating.

The heat producing organ of an aroid is called the spadix. Technically speaking, a spadix is a spike of minute flowers closely arranged around a fleshy axis. All aroid inflorescences have one and they come in a wide variety of shapes, colors, and textures. To produce heat, the spadix is hooked up to a massive underground energy reserve largely in the form of carbohydrates or sugars. The process of turning these sugars into heat is rather complex and surprisingly animal-like.

Cross section of a typical aroid inflorescence with half of the protective spathe removed. The spadix is situated in the middle with a rings of protective hairs (top), male flowers (middle), and female flowers (bottom).

Cross section of a typical aroid inflorescence with half of the protective spathe removed. The spadix is situated in the middle with a rings of protective hairs (top), male flowers (middle), and female flowers (bottom).

It all starts with a compound we are rather familiar with - salicylic acid - as it is the main ingredient in Aspirin. In aroids, however, salicylic acid acts as a hormone whose job it is to initiate both the heating process as well as the production of floral scents. It signals the mitochondria packed inside a ring of sterile flowers located at the base of the spadix to change their metabolic pathway.

In lieu of their normal metabolic pathway, which ends in the production of ATP, the mitochondria switch over to a pathway called the "Alternative Oxidase Metabolic Pathway." When this happens, the mitochondria start burning sugars using oxygen as a fuel source. This form of respiration produces heat.

Thermal imaging of the inflorescence of  Arum maculatum .

Thermal imaging of the inflorescence of Arum maculatum.

As you can imagine, this can be a costly process for plants to undergo. A lot of energy is consumed as the inflorescence heats up. Nonetheless, some aroids can maintain this costly level of respiration intermittently for weeks on end. Take the charismatic skunk cabbage (Symplocarpus foetidus) for example. Its spadix can reach temperatures of upwards of 45 °F (7 °C) on and and off for as long as two weeks. Even more incredible, the plant is able to do this despite freezing ambient temperatures, literally melting its way through layers of snow.

For some aroids, however, carbohydrates just don't cut it. Species like the Brazilian Philodendron bipinnatifidum produce a staggering amount of floral heat and to do so requires a different fuel source - fat. Fats are not a common component of plant metabolisms. Plants simply have less energy requirements than most animals. Still, this wonderful aroid has converged on a fat-burning metabolic pathway that puts many animals to shame. 

The inflorescence of  Philodendron bipinnatifidum  can reach temps as high as 115 °F (46 °C)

The inflorescence of Philodendron bipinnatifidum can reach temps as high as 115 °F (46 °C)

P. bipinnatifidum stores lots of fat in sterile male flowers that are situated between the fertile male and female flowers near the base of the spadix. As soon as the protective spathe opens, the spadix bursts into metabolic action. As the sun starts to set and P. bipinnatifidum's scarab beetle pollinators begin to wake up, heat production starts to hit a crescendo. For about 20 to 40 minutes, the inflorescence of P. bipinnatifidum reaches temperatures as high as 95 °F (35 °C) with one record breaker maxing out at 115 °F (46 °C)! Amazingly, this process is repeated again the following night.

It goes without saying that burning fat at a rate fast enough to reach such temperatures requires a lot of oxygen. Amazingly, for the two nights it is in bloom, the P. bipinnatifidum inflorescence consumes oxygen at a rate comparable to that of a flying hummingbird, which are some of the most metabolically active animals on Earth.

The world's largest inflorescence belongs to the titan arum ( Amorphophallus titanum ) and it too produces heat.

The world's largest inflorescence belongs to the titan arum (Amorphophallus titanum) and it too produces heat.

Again, why these plants go through the effort of heating their reproductive structures is still a bit of a mystery. For most, heat likely plays a role in helping to volatilize floral scents. Anyone that has spent time around blooming aroids knows that this plant family produces a wide range of odors from sweet and spicy to downright offensive. By warming these compounds, the plant may be helping to lure in pollinators from a greater distance away. It is also thought that the heat may be an attractant in and of itself. This is especially true for temperate species like the aforementioned skunk cabbage, which frequently bloom during colder months of the year. Likely both play a role to one degree or another throughout the aroid family.

What we can say is that the process of plant thermogenesis is absolutely fascinating and well worth deeper investigation. We still have much to learn about this charismatic group of plants.

LEARN MORE ABOUT AROID POLLINATION HERE

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

Further Reading: [1] [2] [3] [4] [5] [6] [7] [8]

 

The Intriguing Pollination of a Central American Anthurium

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As an avid gardener of both indoors and out, there are few better experiences than getting to see familiar plants growing in the wild for the first time. That experience is made all the better when you find out new and interesting facts about their ecology. On a recent trip to Costa Rica, I was introduced to a wide variety of Anthurium species. I marveled at how amazing these plants look in situ and was taken aback to learn that many produce flowers with intoxicating aromas.

I was also extremely fortunate to be in the presence of some aroid experts during this trip and their knowledge fueled my interest in getting up close and personal with what little time I had with these plants. They were able to ID the plants and introduce me to their biology. One species in particular has been the subject of interest in an ongoing pollination study that has proven to be unique.

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The plant in question is known scientifically as Anthurium acutifolium and it is rather charming once you get to know it. It is a terrestrial plant with relatively large leaves for its overall size. Its range includes portions of lowland Costa Rica and Panama. Its flowers are typical of what one would expect out of this family. They are fused into a type of inflorescence known as a spadix and can range in color from white to green and occasionally red. If you are lucky to visit the spadix between roughly 8:00 AM and 12:30 PM, you may notice a rich scent that, to me, is impossible to describe in words.

It's this scent that sets the stage for pollination in this species. During some down time, University of Vienna grad student Florian Etl discovered that the spadix of A. acutifolium was getting a lot of attention from a particular species of small bee. Closer inspection revealed that they were all males of a species of oil-collecting bee known as Paratetrapedia chocoensis. Now, the females of these oil collecting bees are well known in the pollination literature. They visit flowers that secrete special oils that the females then use to build nests and feed their young. This is why the attention from male bees was so intriguing.

A: A male  P. chocoensis  bee approaching a scented spadix of an inflorescence of  A. acutifolium . B: The abdominal mopping behavior of male  P. chocoensis  oil bees on a spadix. C: Ventral side of the abdomen of a male  P.chocoensis  covered with pollen. D: A male  P. chocoensis  bee on a spadix of an inflorescence of  A. acutifolium , touching the pollen shedding anthers. E: Pubescent region pressed on the surface of  A. acutifolium  during the mopping behavior. F: A scented inflorescence of  A. acutifolium  with three male  P. chocoensis  individuals. G: Image of the abdomen of a male  P.chocensis  in lateral view showing the conspicuous pubescent region. ( SOURCE )

A: A male P. chocoensis bee approaching a scented spadix of an inflorescence of A. acutifolium. B: The abdominal mopping behavior of male P. chocoensis oil bees on a spadix. C: Ventral side of the abdomen of a male P.chocoensis covered with pollen. D: A male P. chocoensis bee on a spadix of an inflorescence of A. acutifolium, touching the pollen shedding anthers. E: Pubescent region pressed on the surface of A. acutifolium during the mopping behavior. F: A scented inflorescence of A. acutifolium with three male P. chocoensis individuals. G: Image of the abdomen of a male P.chocensis in lateral view showing the conspicuous pubescent region. (SOURCE)

Males would land on the spadix and begin rubbing the bottom of their abdomen along its surface. In doing so, they inevitably picked up and deposited pollen. To date, such behavior was unknown among male oil bees. What exactly were these male bees up to?

As it turns out, the males were collecting fragrances. Close inspection of their morphology revealed that each male has a small patch of dense hairs underneath their abdomen. The males are definitely not after fatty oils or nectar as A. acutifolium does not secrete either of these substances. Instead, it would appear that the male oil bees are there to collect scent, which is mopped up by that dense patch of hairs. Even more remarkable is the fact that in order to properly collect these fragrance compounds, the bees are likely using solvents that they have collected from other flowering plant species around the forest.

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What they are doing with these scent compounds remains a mystery but some potential clues lie in another scent/pollination system. Male orchid bees perform similar scent-collecting activities in order to procure unique scent bouquets. Though the exact function of their scent collecting is not known either, we do know that these scents are used in the process of finding and procuring mates. It is likely that these male oil bees are using them in a similar way.

Taken together, these data suggest that a very specific pollination syndrome involving A. acutifolium and male oil bees has evolved in Central American forests. No other insects were observed visiting the flowers of A. acutifolium and the scents only ever attracted males of these specific oil bees during the hours in which the spadix was actively producing the compounds. This is a remarkable pollination syndrome and one that encourages us to start looking elsewhere in the forest. This, my friends, is why there is no substitute for simply taking the time to observe nature. We must take the time to get outside and poke around because we stand to miss out on so much of what makes our world tick and without such knowledge, we risk losing so much. 

Photo Credits: Florian Etl [1]

Further Reading: [1]

This Isn't Even My Final Form! A Pothos Story

Pothos might be one of the most widely cultivated plants in modern history. These vining aroids are so common that I don't think I can name a single person in my life that hasn't had one in their house at some point or another. Renowned for their hardy disposition and ability to handle extremely low light conditions, they have become famous the world over. They are so common that it is all too easy to forget that they have a wild origin. What's more, few of us ever get to see a mature specimen. The plants living in our homes and offices are mere juveniles, struggling to hang on as they search for a canopy that isn't there.

Trying to find information on the progenitors of these ubiquitous houseplants can be a bit confusing. To do so, one must figure out which species they are talking about. Without a proper scientific name, it is nearly impossible to know which plant to refer to. Common names aside, pothos have also undergone a lot of taxonomic revisions since their introduction to the scientific community. Also, what was thought to be a single species is actually a couple.

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To start with, the plants you have growing in your home are no longer considered Pothos. The genus Pothos seemed to be a dumping ground for a lot of nondescript aroid vines throughout the last century. Many species were placed there until proper materials were thoroughly scrutinized. Today, what we know as a "Pothos" has been moved into the genus Epipremnum. This revision did not put all controversies to rest, however, as the morphological changes these plants go through as they age can make things quite tricky.

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As I mentioned, the plants we keep in our homes are still in their juvenile form. Like all plants, these vines start out small. When they find a solid structure in a decent location, they make their bid for the canopy. Up in a tree in reach of life giving sunlight, these vines really hit their stride. They quickly grow their own version of a canopy that consists of massive leaves nearing 2 feet in length! This is when these plants begin to flower. 

As is typical for the family, the inflorescence consists of a spadix covered by a leafy spathe. The spadix itself is covered in minute flowers and these are the key to properly identifying species. When pothos first made its way into the hands of botanists, all they had to go on were the small, juvenile leaves. This is why their taxonomy had been such a mess for so long. Materials obtained in 1880 were originally named Pothos aureus. It was then moved into the genus Scindapsus in 1908.

Controversy surrounding a proper generic placement continued throughout the 1900's. Then, in the early 1960's, an aroid expert was finally able to get their hands on an inflorescence. By 1964, it was established that these plants did indeed belong in the genus Epipremnum. Sadly, confusion did not end there. The plasticity in forms and colors these vines exhibit left many confusing a handful of species within the group. At various times since the late 1960's, E. aureum and E. pinnatum have been considered two forms of the same species as well as two distinct species. The latest evidence I am aware of is that these two vines are in fact distinct enough to warrant species status. 

The plant we most often encounter is E. aureum. Its long history of following humans wherever they go has led to it becoming an aggressive invader throughout many regions of the world. It is considered a noxious weed in places like Australia, Southeast Asia, India, Pakistan, and Hawai'i (just to name a few). It does so well in these places that it has been a little difficult to figure out where these plants originated. Thanks to some solid detective work, E. aureum is now believed to be native to Mo'orea Island off the west coast of French Polynesia. 

Epipremnum pinnatum  is similar until you see an adult plant

Epipremnum pinnatum is similar until you see an adult plant

It is unlikely that most folks have what it takes to grow this species to its full potential in their home. They are simply too large and require ample sunlight, nutrients, and humidity to hit their stride. Nonetheless there is something to be said for the familiarity we have with these plants. They have managed to enthrall us just enough to be a fixture in so many homes, offices, and shopping centers. It has also helped them conquer far more than the tiny Pacific island on which they evolved. Becoming an invasive species always seems to have a strong human element and this aroid is the perfect example.

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

Further Reading: [1] [2] [3] 

 

Unlikely Allies

On the Balearic Islands of Spain, an interesting relationship has developed between a plant and an animal. What's more, this relationship seems to have developed relatively recently in the history of these two species. The players in this story are the dead horse arum (Helicodiceros muscivorus) and an unsuspecting lizard known as Lilford’s wall lizard (Podarcis lilfordi).

Podarcis lilfordi is a lot like other fence lizards. They spend their days basking in the sun’s warmth and hunting for insect prey. They also have a tendency to feed on nectar and pollen, making them important pollinators of a handful of plant species around the island. For the dead horse arum, however, its not about pollination.

Like most members of its family, the dead horse arum relies on trickery for sex. As its common name suggests, the dead horse arum both looks and smells like rotting meat. Unsuspecting flies looking for a meal and a place to lay their eggs find the dead horse arum quite attractive in this regard. The plant even steps up its game a bit by producing its own heat. This helps volatilize its smell as well as to make it a cozy place worth investigating. Studies have found that during the peak flowering period, the inflorescence can be upwards of 24 °C (50 °F) warmer than its surroundings.

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As one would expect, this has caught the attention of the cold blooded lizards. Enticed by the heat source, lizards basking on the spathe quickly realize that the plant is also a great place to hunt. Flies attracted to and trapped by the flowers make an easy meal. On the surface this would seem counterproductive for the dead horse arum. What good is an animal hanging around that eats its pollinators?

The relationship doesn't end here though. At some point in recent history, a handful of lizards figured out that the seeds of the dead horse arum also make a great meal. This behavior quickly spread through the population to the point that Podarcis lilfordi regularly break open the seed heads and consume the fleshy berries within. Here's the catch, seeds that have passed through a lizards gut are twice as likely to germinate.

Researchers have been studying this interaction since 1999. Since then, the dead horse arum has gone from being relatively rare on the island (~5,000 individuals per hectare) to a density of roughly 30,000 individuals per hectare during the 6 year span of the study! Even though the lizards eat their pollinators, the dead horse arums of Aire Island have nonetheless benefited from interactions with their cold blooded companions.

Sadly, this novel relationship may not last too long. The introduction of cats and rats to the islands has drastically reduced the population of these lizards to the point that the IUCN has listed them as an endangered species. Research will be needed to see if the dead horse arum follows in their wake.

Photo Credit: [1] [2]

Further Reading: [1] [2]