Some Magnolia Flowers Have Built-In Heaters

Magnolia denudata. Photo by 阿橋 HQ licensed under CC BY-SA 2.0

There are a lot of reasons to like magnolias and floral thermogenesis is one of them. That’s right, the flowers of a surprising amount of magnolia species produce their own heat! Although much more work is needed to understand the mechanisms involved in heat generation in these trees, research suggests that it all centers on pollination.

Magnolias have a deep evolutionary history, having arose on this planet some 95+ million years ago. Earth was a very different place back then. For one, familiar insect pollinators like bees had not evolved yet. As such, the basic anatomy of magnolia flowers was in place long before bees could work as a selective pressure in pollination. What were abundant back then were beetles and it is thought that throughout their history, beetles have served as the dominant pollinators for most species. Indeed, even today, beetles dominate the magnolia pollination scene.

Magnolia sprengeri. Photo by Aleš Smrdel licensed under CC BY-NC 2.0

Beetles are generally not visiting flowers for nectar. They are instead after the protein-rich pollen within each anther. It seems that when the anthers are mature, beetles are very willing to spend time munching away within each flower, however, keeping their attention during the female phase of the flower is a bit trickier. Because there are no rewards for visiting a magnolia flower during its female phase, evolution has provided some species with an interesting trick. This is where heat comes in.

Though it varies from species to species, thermogenic magnolias produce combinations of scented oils that various beetles species find irresistible. That is, if they can pick up the odor against the backdrop of all the other enticing scents a forest has to offer. By observing floral development in species like Magnolia sprengeri, researchers have found that as the flowers heat up, the scented oils produced by the flower begin to volatilize. In doing so, the scent is dispersed over a much greater area than it would be without heat.

Magnolia tamaulipana. Photo by James Gaither licensed under CC BY-NC-ND 2.0

Unlike some other thermogenic plants, heat production in magnolia flowers doesn’t appear to be constant. Instead, flowers experience periodic bursts of heat that can see them reaching temperatures as high as 5°C warmer than ambient temperatures. These peaks in heat production just to happen to coincide with the receptivity of male and female organs. Also, only half of the process is considered an “honest signal” to beetles. During the male phase, the beetles will find plenty of pollen to eat. However, during the female phase, the scent belies the fact that beetles will find no reward at all. This has led to the conclusion that the non-rewarding female phase of the magnolia flower is essentially mimicking the rewarding male phase in order to ensure some cross pollination without wasting any energy on additional rewards.

The timing of heat production also changes depending on the species of beetle and their feeding habits. For species like the aforementioned M. sprengeri, which is pollinated by beetles that are active during the day, heat and scent production only occur when the sun is up. Alternatively, for species like M. tamaulipana whose beetle pollinators are nocturnal, heat and scent production only occur at night. Researchers also think that seasonal climate plays a role as well, suggesting that heat itself may be its own form of pollinator reward in some species. Many of the thermogenic magnolias bloom in the early spring when temperatures are relatively low. It is likely that, aside from pollen, beetles may also be seeking a warm spot to rest.

Personally, I was surprised to learn just how many different magnolias are capable of producing heat in their flowers. When I first learned of this phenomenon, I thought it was unique to M. sprengeri but I was wrong. We still have a lot to learn about this process but research like this just goes to show you that even familiar genera can hold many surprises for those curious enough to seek them out.

Further Reading: [1] [2] [3] [4] [5]

How Aroids Turn Up the Heat

Photo by Jörg Hempel licensed under CC BY-SA 2.0

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). Photo by Kristian Peters -- Fabelfroh licensed under CC BY-SA 3.0

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.

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). Photo by Tekwani licensed under CC BY-SA 3.0

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. Photo by Fbianh licensed under CC0 1.0

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]

Do Yeasts Aid Pollination For the Stinking Hellebore?

Photo by Mark Gurney licensed under CC BY-NC-SA 2.0

Whether they are growing in their native habitat or in some far away garden, Hellebores are some of the earliest plants to bloom in the spring. Hellebore flowers can often be seen blooming long before the snow has melted away. All early blooming plant species are faced with the challenge of attracting pollinators. Though the competition for insect attention is minimal among these early bloomers, only the hardiest insects are out and about on cold, dreary days. It stands to reason then that anything that can entice a potential pollinator would be of great benefit for a plant.

That is why the presence of yeast in the nectar of at least one species of Hellebore has attracted the attention of scientists. The species in question is known scientifically as Helleborus foetidus. The lack of appeal in its binomial is nothing compared to its various common names. One can often find H. foetidus for sale under names like the "stinking hellebore" or worse, "dungwort." All of these have to do with the unpleasant aroma given off by its flowers and bruised foliage. Surprisingly, that is not the topic of this post.

Photo by Bernd Haynold licensed under CC BY-SA 3.0

What is more intriguing about the flowers of H. foetidus is that the nectar produced by its smelly green flowers harbors dense colonies of yeast. Yeasts are everywhere on this planet and despite their economic importance, little is known about how they function in nature. For instance, what the heck are these yeast colonies doing in the nectar of this odd Hellebore?

To test this, two researchers from the Spanish National Research Council manipulated yeast colonies within the flowers to see what might be happening. It turns out, yeast in the nectar of H. foetidus actually warms the flowers. As the yeast feed on the sugars within the nectar, their metabolic activity can raise the temperature of the flowers upwards of 2 °C above the ambient. As far as we know, the only other ways in which floral heating has been achieved is either via specific metabolic processes within the floral tissues or by direct heating from the sun. 

In heating the flowers, these yeast colonies may be having serious impacts on the reproductive success of H. foetidus. For starters, these plants are most at home under the forest canopies of central and western Europe. What's more, many populations find themselves growing in the dense shade of evergreens. This completely rules out the ability to utilize solar energy to heat blooms. Additionally, floral heat can mean more visits by potential pollinators. Experiments have shown that bees preferentially visit flowers that are slightly warmer than ambient temperatures. Even the flowers themselves can benefit from that heat. Warmer flowers have higher pollination rates and better seed set.

Bombus terrestris was one of the most common floral visitors of Helleborus foetidus. Photo by Vera Buhl licensed under CC BY-SA 3.0

Yeast colonies also have their downsides. The heat generated by the yeast comes from the digestion of sugars. Indeed, nectar housing yeast colonies had drastically reduced sugar loads than nectar without yeast. This has the potential to offset many of the benefits of floral warming in large part because bees are good at discriminating. Bees are visiting these blooms as a food source and by diminishing the sugar content of the nectar, the yeast may be turning bees off to this potential source. The question then becomes, do bees prefer heat over sugar-rich food? The authors think there might be a trade-off, with bees preferring heated flowers on colder days and sugar-rich flowers on warmer days.

Helleborus foetidus flowering before the snow has had a chance to melt!

Though the authors found evidence for heating, they did not test for pollinator preference. All we know at this point is that yeast in the nectar significantly warms H. foetidus flowers. Since this piece was originally published, more attention has been paid to the benefits of the heat generated from yeast. Interestingly, researchers found that pollen tube formation was higher for H. foetidus flowers that experienced heat earlier in the season but not for those that experienced heat later on. This response, however, was not due to the warming directly. Instead, it had more to do with bee preference.

As it turns out, bumblebees do in fact prefer to visit heated flowers but their preference is limited to the early periods of flowering when ambient temperatures are still quite low. More bumblebees visiting heated flowers in the early spring equated to more pollen being deposited on the stigma, which in turn led to an increase in pollen tube formation and higher seed set. Later on in the season, when ambient temperatures increased a bit, this positive effect dropped off as bees apparently spent more time foraging elsewhere.

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

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