Floral Pigments in a Changing World

Photo by moggafogga licensed under CC BY-NC-ND 2.0

Flowers paint the world in a dazzling array of colors. Some of these we can see and others we cannot. Many plants paint their blooms in special pigments that absorb ultraviolet light, revealing intriguing patterns to pollinators like bees and even some birds that can see well into the UV part of the electromagnetic spectrum. UV absorbing pigments do more than attract pollinators. They can also protect sensitive reproductive organs from UV radiation. By studying these pigments, scientists are finding that many different plants are changing their floral displays in response to changes in their environment.

Growing up I heard a lot about the hole in the ozone layer. Prior to the 1980’s humans were pumping massive quantities of ozone-depleting chemicals such as halocarbon refrigerants, solvents, and chlorofluorocarbons (CFCs) into the atmosphere, creating a massive hole in the ozone layer. Though ozone depletion has improved markedly thanks to regulations placed on these chemicals, it doesn’t mean that life has not had to adapt. As you may remember from your grade school science class, Earth’s ozone layer helps protect life from the damaging effects of ultraviolet radiation. UV radiation damages sensitive biological molecules like DNA so it is in any organisms best interest to minimize its impacts.

UV absorbing pigments in floral tissues can do just that. In addition to attracting pollinators, these pigments act as a sort of sun screen, reducing the likelihood of damaging mutations. By studying 1,238 herbarium specimens collected between 1941 and 2017 representing 42 different species, scientists discovered a startling change in the amount of UV pigments produced in their flowers.

Exemplary images for a species with anthers exposed to ambient conditions, Potentilla crantzii (A–C) and a species with anthers protected by floral tissue Mimulus guttatus (D–F). Darker petal areas possess UV-absorbing compounds whereas lighter areas are UV reflective and lack UV-absorbing compounds. (B) and (E) display a reduced area of UV-absorbing pigmentation on petals compared to (C) and (F). Arrows in (E) and (F) highlight differences in pigment distribution on the lower petal lobe of M. guttatus. [SOURCE]

Across North America, Europe, and Australia, the amount of UV pigments produced in the flowers tended to increase by an average of 2% per year from 1941 to 2017. These increases in UV pigments occurred in tandem with decreases in the ozone layer. It would appear that, to protect their reproductive organs from harmful UV rays, many plants were increasing these protective pigments.

However, changes in UV pigments were not uniform across all the species they examined. Plants that produce saucer or cup-shaped flowers experienced the greatest increases in UV pigments. This makes complete sense as this sort of floral morphology exposes the reproductive organs directly to the sun’s rays. The pattern reversed when scientists examined flowers whose petals enclose the reproductive organs such as those seen in bladderworts (Utricularia spp.). UV pigments in flowers that conceal their reproductive organs actually decreased over this time period.

The reason for this comes down to a trade off inherent in UV pigments. Absorbing UV radiation is a great way to reduce its impact on sensitive tissues but it also leads to increased temperatures. For plants that enclose their reproductive organs within their petals, this can lead to overheating. Heat can also be very damaging to floral structures so it makes complete sense that species with this type of floral morphology would demonstrate the opposite pattern. By reducing the amount of UV absorbing pigments in their flowers, plants like bladderworts are able to minimize the effect of increased radiation and temperatures that occurred over this time period.

How changes in floral pigments are affecting pollination rates for these plants is another story entirely. Because UV pigments also help attract certain pollinators, there is always a chance that the appearance of some of these flowers may also be changing over time. Now that we know this is occurring across a wide range of unrelated plants, research can now be aimed at tackling questions like this.

Photo Credits: [1] [2]

Further Reading: [1]

Buzzing Bees Make Evening Primrose Flowers Sweeter

Photo by Guy Haimovitch licensed under CC BY-ND 2.0.

Plants, like all living organisms, must be able to sense and respond to their environment. The more we look at these sessile organisms, the more we realize that plants are far from static in their day to day lives. Recent evidence even suggests that some plants may be able to “hear” their pollinators and react accordingly.

I place the word “hear” in quotes because I want to make sure that we are not talking about hearing in an animalistic sense. Plants do not have ears, a nervous system, or anything like a central processing unit to make sense of such stimuli. What they do have are mechanoreceptors that can sense vibrations and those are what are likely at work in this example.

The beach evening primrose (Oenothera drummondii) is native to southeastern North America. It is pollinated by bees during the day and by moths at night. Like most members of its genus, O. drummondii produces relatively large, showy flowers. That doesn’t mean it steals all of the attention though. Competition for pollinators can be stiff among flowering plants. To sweeten the deal a bit, O. drummondii also produces a fair amount of nectar.

Nectar is costly for plants to produce and maintain. Not only does it take water and carbohydrates away from the rest of the plant, it also puts the reproductive structures at risk of degradation by microbes feeding on sugars as well as nectar thieves who end up drinking the nectar without pollinating the flower. It stands to reason that a plant that can modulate the quality of its nectar reward in response to pollinator availability could potentially increase its fitness. If the plant doesn’t always have to present sugar-rich nectar then why bother? It appears that selective nectar production is exactly the strategy O. drummondii employs.

Photo by Yu-Ju Chang licensed under CC BY-ND 2.0.

Researchers have discovered that individual O. drummondii flowers can rapidly increase the sugar content of their nectar after being exposed to the sound of a visiting bee. Within 3 minutes of being exposed to playbacks of bee wings, the flowers of O. dummondii increased the sugar content of their nectar by 20%. What’s more, flowers that had sensed the vibrations and increased their sugar content were more likely to be visited by bees. This is because bees are really good at sensing the sugar content of nectar.

This is pretty remarkable. Not only does this enable the plant to respond to the availability of pollinators and reduce the chances of nectar spoilage and theft, it significantly increases their chances of pollination. The fact that the response is so rapid (~3 mins) likely stems from the foraging habits of bees, who prefer to limit the amount of time between floral visits. Thus, the faster the plant can respond, the more likely that bees are willing to stick around and visit more flowers.

In terms of a mechanism, researchers believe the flower itself is the main sensory organ involved in the response. As mentioned, plants do produce mechanoreceptor proteins, which can sense physical vibrations. The presence of these proteins within the petals likely plays a role in sensing bee vibrations. Moreover, the bowl-shape of the flower itself may be under some selective pressures that favor the ability of the flower to sense its pollinators. More work is needed to better understand exactly how the signal pathways play out. Also, the question remains as to how wide spread this phenomenon is and how it differs between different plants and floral shapes.

Photo Credits: [1] [2]

Further Reading: [1]

Deer Skew Jack-in-the-Pulpit Sex Ratios

Photo by Michael Janke licensed under CC BY-ND 2.0.

Deer populations in North America are higher than they have been at any point in history. Their explosion in numbers not only leads to series health issues like starvation and chronic wasting disease, it has also had serious impacts on regional plant diversity. Wherever hungry herds of deer go, plants disappear from the landscape. However, the impacts of deer on plants aren’t limited to species they can eat. Research on Jack-in-the-Pulpit (Arisaema triphyllum) has shown that deer can have plenty of surprising indirect impacts on plants as well.

Though I wouldn’t put anything past a hungry deer, plants like Jack-in-the-Pulpit aren’t usually on the menu for these ungulates. Their leaves, stems, and flowers are chock full of raphide crystals that will burn the mouths and esophagus of most herbivores. Still, this doesn’t mean deer aren’t impacting these plants in other ways. Because deer are congregating in high abundance in our ever-shrinking natural spaces, they are having serious impacts on local growing conditions. Wherever deer herds are at high numbers, forests are experiencing soil compaction, soil erosion, and a disappearance of soil leaf litter (also due in part to invasive earthworms). Thanks to issues like these, plants like Jack-in-the-Pulpit are undergoing some serious changes.

Like many aroids, sex expression in the genus Arisaema is fluid and relies on energy stores. Smaller plants store less energy and tend to only produce male flowers when they bloom. Pollen, after all, is cheap compared to eggs and fruit. Only when a plant has stored enough energy over the years will it begin to produce female flowers in addition to males and only the largest, most robust plants will switch over entirely to female flowers. As you can imagine, the ability of a plant to acquire and store enough energy is dependent on the quality of the habitat in which it grows. This is where deer enter into the equation.

High densities of deer inevitably cause serious declines in habitat quality of plants like Jack-in-the-Pulpit. As leaf litter disappears and soil compaction grows more severe, individual plants have a much harder time storing enough energy each growing season. In places where deer impacts are heaviest, the sex ratios of Jack-in-the-Pulpit populations begin to skew heavily towards males because individual plants must grow much longer before they can store enough energy to produce female flowers. It doesn’t end there either. Not only does it take longer for a plant to begin producing female flowers, individual plants must also reach a much larger size in order to produce female flowers than in areas with fewer deer.

Photo by Charles de Mille-Isles licensed under CC BY-ND 2.0.

As mentioned, seed production takes a lot of energy and any plant that is able to produce viable fruits will have less energy stores going into the next season. This means that even if a plant is able to produce female flowers and successfully set seed, they will have burned through so much energy that they will likely revert right back to producing only male flowers the following year, further skewing the sex ratios of any given population towards males. Interestingly, this often results in more individuals being produced via clonal offshoots. The more clones there are in a population, the less diverse the gene pool of that population becomes.

Without actually eating the plants, deer are having serious impacts on Jack-in-the-Pulpit population dynamics. I am certain that this species isn’t alone either. At least Jack-in-the-Pulpit is somewhat flexible in its reproductive behaviors. Other plants aren’t so lucky. I realize deer are a hot button issue but there is no getting around the fact that our mismanagement of their natural predators, habitat, and numbers are having serious and detrimental impacts on wild spaces and all the species they support.

Photo Credits: [1] [2]

Further Reading: [1]

To grow or to flower, that is the cactus conundrum

Melocactus intortus

Flowers are costly structures for plants to produce. In the flowering plant world, there is always a trade-off between growth and reproduction. Flowers are produced from tiny structures called axillary buds, and many plants can only produce one flush of flowers per bud. Cacti are no exception to this rule and their amazing morphological adaptations to harsh climates has forced them into quite a conundrum when it comes to reproduction.

The axillary buds of cacti are located at the base of their spines in little structures called areoles. This is where the flowers will eventually emerge. However, unlike plants that can produce cheap stems and branches, cacti must produce a whole new chunk of stem or internode before they can produce more axillary buds. Think of it this way, if a cactus wants to produce 10 flowers, it must produce ten internodes to do so. This means producing all of the expensive cortex and epidermis along with it. Their harsh environments have forced most cacti into an extremely tight relationship between growth, water storage, photosynthesis, and flowering that is potentially very limiting from a reproductive standpoint.

Micranthocereus estevesii with lateral cephalium

Amazingly, some cacti have managed to break from this evolutionary relationship and they have done so in a bizarre way. Take a look at all of the cacti pictured here. Each has developed a strange looking structure called a cephalium. Essentially, you can think of the cephalium of a cactus as its “adult” reproductive form whereas the rest of the body consists of non-reproductive, photosynthetic “juvenile” form.

The cephalium is a unique and fascinating structure. It differs from the rest of the cactus body in that it is not photosynthetic. It also produces no chlorophyll and no stomata. In fact, it does not form anything like the epidermis of the rest of the plant. Instead, the cephalium produces dense clusters of short spines and trichomes. Most importantly, it produces tightly packed axillary buds in high abundance. These are the buds that will produce the flowers. The end result is a wacky looking structure that has the ability to produce far more flowers than that of cacti that do not grow a cephalium.

Facheiroa tenebrosa with lateral cephalium

Obviously not all cacti produce cephalia but it is common in genera such as Melocactus, Backebergia, Espostoa, Discocactus, and Facheiroa (this is not a complete list). What the cephalium has done for genera like these is decouple the afore mentioned relationships between growth and reproduction. For a period of time (often many years) following germination, these cacti grow the typical succulent, photosynthetic stems we are accustomed to seeing.

At some point in their development, something triggers these plants to switch to their adult forms. Axillary buds within either lateral or apical meristems switch their growth habit and begin forming the cephalium. It is worth mentioning that no one yet knows what triggers this switch. If the cephalium is produced from axillary buds in the apical meristem like we see in Melocactus, the plant will no longer produce photosynthetic tissues. This represents another major trade-off for these cacti. Such species must rely on the photosynthetic juvenile tissues for all of their photosynthetic needs for the rest of their lives (unless the cephalium is damaged or lost). Backebergia have managed to get around this trade-off by not only growing multiple stems, they will also shed their apical cephalia after a few years, thus re-initiating photosynthetic juvenile growth.

Backebergia militaris with bizarre apical cephalia reminiscent of the bearskin hats of the Queen’s guard.

Things are a bit different for cacti that produce lateral cephalia. Genera such as Espostoa, Facheiroa, and Buiningia are less limited by their cephalia because they are produced along the ribs of the stem, thus leaving the apical meristem free to continue more typical photosynthetic growth. Nonetheless, the process is much the same. Dense clusters of spines, trichomes, and most importantly, axillary buds are produced along the rib, giving each stem a lovely, lopsided appearance.

There are other benefits to growing cephalia in addition to simply being able to produce more flowers. The densely packed spines and trichomes offer the developing flowers and fruits ample protection from both the elements and herbivores. Floral buds are free to develop deep within the interior of the cephalium until they are mature. At that point, the cells will begin to swell with water, pushing the flower outward from the cephalium where it will be exposed to pollinators. As the petals curl back, they offer a safe spot for visiting pollinators that is free from menacing spines. Once pollination has been achieved, the flower wilts and the deeply inferior ovaries are then free to develop within the safety of the cephalium. Once the fruits are mature, they too will begin to swell with water and be pushed out from the cephalium where they will attract potential seed dispersers.

Melocactus violaceus with fruits emerging from the cephalium

I hope that I have convinced you of just how awesome this growth form can be. I will never forget the first time I saw a cactus topped with a cephalium. It was a mature Melocactus growing in a cactus house. Sticking out of the odd “cap” on top was a ring of bright pink fruits. I knew nothing of the structure at that time but it was incredible to see. Now that I know what it is and how it functions, I am all the more appreciative of these cacti.

This post was inspired by the diligent work of Dr. Jim Mauseth. Click here to learn more about cacti.

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

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

The Succulent Passionflowers

Photo by Wendy Cutler licensed by CC BY-SA 2.0

Succulent passionflowers?! It took me a minute to get my head wrapped around the idea. It wasn’t until I saw one in flower that I truly understood. The genus Adenia is found throughout east and west Africa, Southeast Asia, and hits its peak diversity in Madagascar. It comprises approximately 100 species and, as a whole, is poorly understood. Today I would like to introduce you to this bizarre genus within Passifloraceae.

Adenia glauca Photo by Karelj licensed under the GNU Free Documentation License

Adenia is, to date, the second largest genus within the Passionflower family and yet delineating species has been something of a nightmare for botanists over the years. At least some of this confusion lies within the diversity of this odd group. It has been said that few angiosperm lineages surpass Adenia in the diversity of growth forms they exhibit. Though all could be considered succulent to some degree, Adenia runs the gamut from trees to vines, and even tuberous herbs.

Even within individual species, the overall form of these plants can vary widely depending on the conditions under which they have been growing. Their succulent nature and that fact that many species can reach rather large proportions means that herbarium records for this group are scant at best. Many are only known from a single, incomplete collection of a few bits and pieces of plant. Also, juvenile plants often look very different from their adult forms, making timing of the collection crucial for proper analysis.

To complicate matters more, all Adenia are dioecious, meaning that individual plants are either male or female. Male and female flowers of individual species look pretty distinct and differ a bit from what we have come to expect out of the passionflower family. Often collections were made on only a single sex. This is further complicated by the fact that these plants often exhibit very short flowering seasons. Most come into bloom right before the onset of the rainy season and are entirely leafless at that point in time. Because of this, it has been extremely difficult to accurately match flowering collections to vegetative collections. As such, nearly 1/4 of all Adenia species are missing descriptions of either male or female flowers and their fruits.

Female flower of Adenia reticulata. Photo by C. E. Timothy Paine licensed under CC BY-NC 2.0

Male flowers of Adenia digitata. Photo by Joachim Beyenbach licensed under CC BY-SA 3.0

Flowers of Adenia firingalavensis. Photo by voyage-madagascar.org licensed under CC BY 2.0

Fruits of Adenia hondala

Even genetic work has failed to clear up much of the mysteries that surround this group. Some studies suggest that Adenia is sister to all other genera within Passifloraceae whereas others have even suggested it to be nestled neatly within the genus Passiflora. The most recent work hints at a placement among the tribe Passifloreae. If this confuses you, you are certainly not alone. Until a more complete sampling effort is done on Adenia, I think it is safe to say that this genus will be holding onto its taxonomic mysteries for the foreseeable future.

Adenia globosa photo by KENPEI licensed under the GNU Free Documentation License

All Adenia are perennial plants but how they manage this differs from species to species. Some put all of their energy into underground tubers, producing annual stems and leaves that die back each year. Others don’t produce any tubers and instead store all of their water and nutrients within thick stems. This has made at least a handful of species a hit with succulent growers around the world. It is always an interesting sight to see a giant caudiciform trunk or base with bunches of spindly stems spraying out from the top.

Leaves and fruit of Adenia cissampeloides. Photo by International Institute of Tropical Agriculture licensed under CC BY-NC 2.0

Juvenile Adenia glauca. Photo by laurent houmeau licensed under CC BY-SA 2.0

Adenia are also extremely toxic plants. The conditions under which these plants evolved are tough and it appears that this group doesn’t want to take any chances on losing any biomass to herbivores. The main class of compounds they produce are called lectins. These proteins cause myriad issues within animal bodies including rapid cell death, blood clotting, inhibition of protein synthesis, and a disruption of ribosome and DNA function. Needless to say, its in any critters best interest to avoid nibbling on any species of Adenia. Even handling and pruning of these plants merits caution.

Photo by Wendy Cutler licensed under CC BY 2.0

Whether you’re a botanist, taxonomist, gardener, or just curious about plant diversity, Adenia is a wonderful example of just how many unknowns are still out there. Regardless of their taxonomic status, these are fascinating species, each with a wonderful ecology and intriguing evolutionary history. These plants are hardy survivors and a great example of the lengths a genus can go to when presented with new opportunities. Undoubtedly many more species await description but the plants we currently know of are fascinating to say the least.

Adenia pechuelii. Photo by Ewald Schmidt licensed under public domain.

Photo Credits: [1] [2] [3] [4] [5] [6] [7] [8] [9] [10] [11] [12] [13]

Further Reading: [1] [2]

Meet the Fire Lily

Photo by Callan Cohen licensed under CC BY-SA 3.0

The flora of the South African fynbos region is no stranger to fire. Many species have adapted to cope with and even rely on fire to complete their lifecycles. There is one species, however, that takes this to the extreme. It is a tiny member of the Amaryllidaceae aptly named the fire lily (Cyrtanthus ventricosus).

The fire lily is not a big plant by any means. Mature individuals can top out around 9 inches (250 mm) and for most of the year consist of a nothing more than a small cluster of narrow, linear leaves. As the dry months of summer approach, the leaves senesce and the plant more or less disappears until its time to flower. However, unlike other plants in this region that flower more regularly, the fire lily lies in wait for a very specific flowering cue - smoke.

It has been noted that fire lilies only seem to want to reproduce after a fire. No other environmental factor seems to trigger flowering. This has made them quite frustrating for bulb aficionados. Only after a fire burns over the landscape will a scape emerge topped with anywhere from 1 to 12 tubular red flowers.

This dependence on fire for flowering has garnered the attention of a few botanists concerned with conservation of pyrophytic geophytes. Obviously if we care about conserving species like the fire lily, it is extremely important that we understand their reproductive ecology. The question of fire lily blooming is one of triggers. What part of the burning process triggers these plants to bloom?

By experimenting with various burn and smoke treatments, researchers were able to deduce that it wasn’t heat that triggered flowering but rather something in the smoke itself. Though researchers were not able to isolate the exact chemical(s) responsible, at least we now know that fire lilies can be coaxed into flowering using smoke alone. This is a real boon to growers and conservationists alike.

Photo by Callan Cohen licensed under CC BY-SA 3.0

Seeing a population of fire lilies in full bloom must be an incredible sight. Within only a few days of a fire, huge patches of bright red flowers decorate the charred landscape. They are borne on hollow stalks which provide lots of structural integrity while being cheap to produce. The flowers themselves are not scented but they do produce a fair amount of nectar. The bright red inflorescence mainly attracts the Table Mountain pride butterfly as well as sunbirds.

Once flowering is complete, seeds are produced and the plants return to their dormant bulbous state until winter when leaves emerge again. Flowering will not happen again until fire returns to clear the landscape. This strategy may seem inefficient on the part of the plant. Why not attempt to reproduce every year? The answer is competition. By waiting for fire, this tiny plant is able to make a big impact despite being so small. It would be impossible to miss their enticing floral display when all other vegetation has been burned away.

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

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

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]

Mt. Cuba Center Puts Nativars to the Test

By this point, most gardeners will have undoubtedly heard about the importance of using native plants in our landscapes. Though the idea is not new, Doug Tallamy’s landmark publication “Bringing Nature Home” put native plants on the radar for more gardeners than ever. There is no debate that utilizing native plants in our landscapes offers us a chance to bring back some of the biodiversity that was lost when our homes and work places were built. And, at the end of the day, who doesn’t love the sight of a swallowtail butterfly flitting from flower to flower or a pair of warblers nesting in their Viburnum? The rise of native plants in horticulture and landscaping is truly something worth celebrating.

At the same time, however, capitalism is capitalism, and many nurseries are starting to jump on the bandwagon in alarming ways. The rise of native cultivars or “nativars” is troubling to many. Nativars are unique forms, colors, and shapes of our beloved native plants which have been selected and propagated by nurseries and plant breeders. This has led many to denounce the practice of planting nativars as a slap in the face to the concept of native gardening.

Nativars are frequently seen as unnatural mutant versions of their wild counterparts whose use overlooks the whole point of natives in the first place. Take, for instance, the popularity of double flowered nativars. These plants have been selected for an over-production of sepals and petals that can be so dense that they preclude visitation by pollinators. Another example that will be familiar to most are the bright blue hydrangeas that have become to popular. These shrubs have been selected for producing bright, showy flowers that, depending on your soil chemistry, exhibit a stunning blue coloration. The downside here is that all of those flowers are sterile and produce no nectar or pollen for visiting insects.

It would seem that nativars are a slippery slope to yet another sterile landscape incapable of supporting biodiversity. However, anecdotes don’t equal data and that is where places like Mt. Cuba Center come in. Located in northern Delaware, Mt. Cuba is doing something quite amazing for the sake of environmentally friendly landscaping – they are putting plants to the test.

Mt. Cuba has been running trial garden research and experiments on native plants and their nativars for over a decade. The goal of this research is to generate and analyze data in order to help the public make better, more sustainable choices for their yards. Mt. Cuba aims to better understand and quantify the horticultural and ecological value of native plants and related nativars in order to better understand the various ecosystem services these plants provide. In collaboration with academic institutions in the region, popular nativars are established and grown under similar conditions to those experienced in the yards of your average gardener. They are monitored for years to assess their overall health, performance, and ability to support wildlife. Thanks to the help of countless volunteers, these trial gardens paint a holistic picture of each plant and related nativars that is sorely lacking from the gardening lexicon.

This is very exciting research to say the least. The data coming out of the Mt. Cuba trial gardens may both surprise and excite gardeners throughout the mid-Atlantic region of North America. For instance, their latest report looked at some of the most common Phlox varieties on the market. At the top of this list is Garden Phlox (Phlox paniculata). This lovely species is native throughout much of the eastern United States and has become quite a rockstar in the nursery trade. Over 580 cultivars and hybrids have been named to date and no doubt many more will be introduced in the future. Amazingly, many of these Phlox nativars are being developed in the Netherlands. As such, Phlox arriving in regions of the US with vastly different climates often fall victim to novel diseases they never encountered in Europe. What’s more, people often plant these nativars in hopes of attracting butterflies to their garden. Despite their popularity for attracting various lepidopterans, no one has ever tested whether or not the nativars perform as well as their native progenitor.

Phlox paniculata 'Delta Snow'

Starting in 2015, Mt. Cuba began trials on 66 selections and hybrids of Garden Phlox along with 28 other sun-loving types of Phlox. The plants were observed on a regular basis to see which of the nativars experienced the least amount of disease and attracted the most insects. The clear winner of these trails is a nativar known as Phlox paniculata ‘Jeana’. This particular selection was discovered growing along the Harpeth River in Tennessee and is known for having the smallest flowers of any of the Garden Phlox varieties. It also has the reputation for being rather resistant to powdery mildew. Alongside other selections such as Delta Sno’ and David, Jeana really held up to this reputation.

As far as butterflies are concerned, Jeana blew its competition out of the water. Throughout the observation period, Jeana plants received over 530 visits from butterflies whereas the second place selection, Lavelle, received 117. A graduate student at the University of Delaware is studying why exactly the various nativars of Phlox paniculata differ so much in insect visitation. Though they haven’t zeroed in on a single cause at this point, they suggest that the popularity of Jeana might actually have something to do with its small flower size. Perhaps the density of smaller flowers allows butterflies to access more nectar for less effort.

Phlox paniculata ‘Jeana’

Monarda is another genus of North American native plants that has seen an explosion in nativars and hybrids over the last few decades. The popularity of these mints is no surprise to anyone who has spent time around them. Their inflorescence seems to be doing their best impression of a fireworks display, an attribute that isn’t lost on pollinators. These plants are popular with a wide variety of wildlife from solitary bees to voracious hummingbirds. Even after flowering, their seeds provide food for seed-eating birds and many other animals.

As with Garden Phlox, a majority of the commercial selection and hybridization of Monarda occurs in Europe. As a result, resistance to North American plant diseases is not top priority. Many of us have experienced this first hand as our beloved bee balm patch succumbs to aggressive strains of powdery mildew. Though there are many species of Monarda native to North America, most of the plants we encounter are nativars and hybrids of two species – Monarda didyma and Monarda fistulosa.

Monarda fistulosa 'Claire Grace'

Again, Mt. Cuba’s trial gardens put these plants to the test. A total of 40 different Monarda selections were grown, observed, and ranked based on their overall growth and vigor, pollinator attractiveness, and disease resistance. The clear winner of these trials was a naturally-occurring form of M. fistulosa affectionately named ‘Claire Grace.’ Its floral display lasts a total of 3 weeks without waning and managed to attract over 130 visits by butterflies and moths. Though plenty of other insects such as short-tongued bees visited the flowers during the trial period, they are too small to properly access the nectar inside the flower tubes and are therefore not considered effective pollinators.

Another clear winner in terms of pollinators was possibly one of the most stunning Monarda selections in existence – Monarda didyma ‘Jacob Cline’. This tall, red-flowering nativar was a major hit with hummingbirds. During the observation period, Jacob Cline received over 270 visits from these brightly colored birds. Researchers are still trying to figure out why exactly this particular selection was such a hit but they speculate that the large flower size presents ample feeding opportunities for tenacious hummingbirds.

Monarda didyma 'Jacob Cline'

Claire Grace and Jacob Cline also outperformed most of the other selections in terms of disease resistance. Even in the crowded conditions experienced by plants in the trail garden, both selections faired quite well against the dreaded powdery mildew. Though they aren’t completely resistant to it, these and others did not succumb like some selections tend to do. Interestingly enough, most of the other pure species tested in the trial faired quite well against powdery mildew as well. It would appear that Mother Nature better equips these plants than European breeders.

These reports are but two of the many trials that Mt. Cuba has undertaken and there are many, many more on the way. Thanks to the hard work of staff and volunteers, Mt. Cuba is finally putting numbers behind some of our most commonly held assumptions about gardening with native plants and their cultivars. It is impressive to see a place so dedicated to making our landscapes more sustainable and environmentally friendly.

If you would like to find out more about Mt. Cuba’s trial garden as well as download your own copies of the trial garden reports, please make sure to check out https://mtcubacenter.org/research/trial-garden/

Hydatellaceae: The Other Basal Angiosperms

Photo by Kevin Thiele licensed under CC BY 2.0

Though rather obscure to most of the world, the genus Trithuria has enjoyed somewhat of a celebrity status in recent years. A paper published in 2007 lifted this tiny group of minuscule aquatic plants out of their spot in Poales and granted them a place among the basal angiosperm lineage Nymphaeales. This was a huge move for such little plants. 

The genus Trithuria contains 12 species, the majority of which reside in Australia, however, two species, T. inconspicua and T. konkanensis, are native to New Zealand and India. They are all aquatic herbs and their diminutive size and inconspicuous appearance make them easy to miss. For quite some time these odd plants were considered to be a group of highly reduced monocots. Their original placement was in the family Centrolepidaceae. All of that changed in 2007.

Close inspection of Trithuria DNA told a much different story. These were not highly reduced monocots after all. Instead, multiple analyses revealed that Trithuria were actually members of the basal angiosperm lineage Nymphaeales. Together with the water lilies (Nymphaeaceae) and the fanworts (Cabombaceae), these plants are living representatives of some of the early days in flowering plant evolution. 

Of course, DNA analysis cannot stand on its own. The results of the new phylogeny had to be corroborated with anatomical evidence. Indeed, closer inspection of the anatomy of Trithuria revealed that these plants are truly distinct from members of Poales based on a series of features including furrowed pollen grains, inverted ovules, and abundant starchy seed storage tissues. Taken together, all of these lines of evidence warranted the construction of a new family - Hydatellaceae.

The 12 species of Trithuria are rather similar in their habits. Many live a largely submerged aquatic lifestyle in shallow estuarine habitats. As you may have guessed, individual plants look like tiny grass-like rosettes. Their small flower size has lent to some of their taxonomic confusion over the years. What was once thought of as individual flowers were revealed to be clusters or heads of highly reduced individual flowers. 

Reproduction for these plants seems like a tricky affair. Some have speculated that water plays a role but close inspections of at least one species revealed that very little pollen transfer takes place in this way. Wind is probably the most common way in which pollen from one plant finds its way to another, however, the reduced size of these flowers and their annual nature means there isn't much time and pollen to go around. It is likely that most of the 12 species of Trithuria are self-pollinated. This is probably quite useful considering the unpredictable nature of their aquatic habitats. It doesn't take much for these tiny aquatic herbs to establish new populations. In total, Trithuria stands as living proof that big things often come in small packages. 

Photo Credits: [1]

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

The Extraordinary Catasetum Orchids

Male Catasetum osculatum. Photo by Orchi licensed under CC BY-SA 3.0

Orchids, in general, have perfect flowers in that they contain both male and female organs. However, in a family this large, exceptions to the rules are always around the corner. Take, for instance, orchids in the genus Catasetum. With something like 166 described species, this genus is interesting in that individual plants produce either male or female flowers. What's more, the floral morphology of the individual sexes are so distinctly different from one another that some were originally described as distinct species. 

Female Catasetum osculatum. Photo by Valdison Aparecido Gil licensed under CC BY-SA 4.0

In fact, it was Charles Darwin himself that first worked out that plants of the different sexes were indeed the same species. The genus Catasetum enthralled Darwin and he was able to procure many specimens from his friends for study. Resolving the distinct floral morphology wasn't his only contribution to our understanding of these orchids, he also described their unique pollination mechanism. The details of this process are so bizarre that Darwin was actually ridiculed by some scientists of the time. Yet again, Darwin was right. 

Catasetum longifolium. Photo by Maarten Sepp licensed under CC BY-SA 4.0

If having individual male and female plants wasn't strange enough for these orchids, the mechanism by which pollination is achieved is quite explosive... literally. 

Catasetum orchids are pollinated by large Euglossine bees. Attracted to the male flowers by their alluring scent, the bees land on the lip and begin to probe the flower. Above the lip sits two hair-like structures. When a bee contacts these hairs, a structure containing sacs of pollen called a pollinia is launched downwards towards the bee. A sticky pad at the base ensures that once it hits the bee, it sticks tight. 

Male Catasetum flower in action. Taken from BBC's Kingdom of Plants.

Bees soon learn that the male flowers are rather unpleasant places to visit so they set off in search of a meal that doesn't pummel them. This is quite possibly why the flowers of the individual sexes look so different from one another. As the bees visit the female flowers, the pollen sacs on their back slip into a perfect groove and thus pollination is achieved. 

The uniqueness of this reproductive strategy has earned the Catasetum orchids a place in the spotlight among botanists and horticulturists alike. It begs the question, how is sex determined in these orchids? Is it genetic or are there certain environmental factors that push the plant in either direction? As it turns out, light availability may be one of the most important cues for sex determination in Catasetum. 

Photo by faatura licensed under CC BY-NC-ND 2.0

A paper published back in 1991 found that there were interesting patterns of sex ratios for at least one species of Catasetum. Female plants were found more often in younger forests whereas the ratios approached an even 1:1 in older forests. What the researchers found was that plants are more likely to produce female flowers under open canopies and male flowers under closed canopies. In this instance, younger forests are more open than older, more mature forests, which may explain the patterns they found in the wild. It is possible that, because seed production is such a costly endeavor for plants, individuals with access to more light are better suited for female status. 

Catasetum macrocarpum. Photo by maarten sepp licensed under CC BY-SA 2.0

Aside from their odd reproductive habits, the ecology of these plants is also quite fascinating. Found throughout the New World tropics, Catasetum orchids live as epiphytes on the limbs and trunks of trees. Living in the canopy like this can be stressful and these orchids have evolved accordingly. For starters, they are deciduous. Most of the habitats in which they occur experience a dry season. As the rains fade, the plants will drop their leaves, leaving behind a dense cluster of green pseudobulbs. These bulbous structures serve as energy and water stores that will fuel growth as soon as the rains return. 

Catasetum silvestre in situ. Photo by Antonio Garces licensed under CC BY-NC-ND 2.0

The canopy can also be low in vital nutrients like nitrogen and phosphorus. As is true for all orchids, Catasetum rely on an intimate partnership with special mychorrizal fungi to supplement these ingredients. Such partnerships are vital for germination and growth. However, the fungi that they partner with feed on dead wood, which is low in nitrogen. This has led to yet another intricate and highly specialized relationship for at least some members of this orchid genus. 

Photo by faatura licensed under CC BY-NC-ND 2.0

Mature Catasetum are often found growing right out of arboreal ant nests. Those that aren't will often house entire ant colonies inside their hollowed out pseudobulbs. This will sometimes even happen in a greenhouse setting, much to the chagrin of many orchid growers. The partnership with ants is twofold. In setting up shop within the orchid or around its roots, the ants provide the plant with a vital source of nitrogen in the form of feces and other waste products. At the same time, the ants will viciously attack anything that may threaten their nest. In doing so, they keep many potential herbivores at bay.  

Female Catasetum planiceps. Photo by sunoochi licensed under CC BY 2.0

To look upon a flowering Catasetum is quite remarkable. They truly are marvels of evolution and living proof that there seems to be no end to what orchids have done in the name of survival. Luckily for most of us, one doesn't have to travel to the jungles and scale a tree just to see one of these orchids up close. Their success in the horticultural trade means that most botanical gardens house at least a species or two. If and when you do encounter a Catasetum, do yourself a favor and take time to admire it in all of its glory. You will be happy that you did. 

Photo Credits: [1] [2] [3] [4] [5] [6] [7] [8] [9] 

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

In Search of Fall Flowering Gentians

Fall is bittersweet for avid botanizers but if you look around you, there are still flowers to be seen! Join us as we go in search of some of my favorite fall flowering species of Gentian.

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The Hidden Anatomy of Grass Flowers

Grass flowers have their own unique beauty. Examine them with a hand lens and a whole new world of angiosperm diversity suddenly opens up. Unlike other flowering plants, their charm lies not in showy sepals or petals, but in an intricacy centered around the utilization of wind for pollination. However, such floral organs are not lacking. Grass flowers do in fact produce a perianth, the function of which has been highly modified.

To see what I am referring to, you need to do some dissection under a scope. Pull off a flower and peel away the sheaths (the palea and lemma) that cover it. Inside you will see an ovary complete with feathery stigmas as well as the anthers. At the base of the ovary sits a pair of scales called lodicules. These lodicules are thought to be the rudimentary remains of the perianth. They certainly don't resemble sepals or petals but that is because the function of these structures is not to attract pollinators. They assist in pollination in another way.

Photo by Matt Lavin CC BY-SA 2.0

When grass flowers are ready for reproduction, the lodicules begin to swell. This swelling serves to push apart the rigid palea and lemma that protected the flowering parts as they developed. Once apart, the anthers and stigma are free to emerge and let wind do the dirty work for them. Lodicules differ quite a bit from species to species in their size, shape, and overall appearance. Much of this is likely tied to the overall structure in grass flowers.

Photo Credits: [1] [2]

Further Reading: [1]

Mighty Magnolias

Magnolias are one of those trees that even the non-botanically minded among us will easily recognize. They are one of the more popular plant groups grown as ornamentals and their symbolism throughout human history is quite interesting. But, for all this attention, few may realize how special magnolias really are. Did you know they they are one of the most ancient flowering plant lineages in existence?

Magnolias first came on to the scene somewhere around 95 million years ago. Although they are not representative of what the earliest flowering plants may have looked like, they do offer us some interesting insights into the evolution of flowers. To start with, the flower bud is enclosed in bracts (modified leaves) instead of more differentiated sepals. The "petals" themselves are not actually petals but tepals, which are also undifferentiated. The most striking aspect of magnolia flower morphology is in the actual reproductive structures themselves.

Magnolias evolved before there were bees. Because of this, the basic structure that makes them unique was in place long before bees could work as a selective pressure in pollination. Beetles are the real pollinators of magnolia flowers. The flowers have a hardened carpel to avoid damage by their gnawing mandibles as the feed. The beetles are after the protein-rich pollen. Because the beetles are interesting in pollen and pollen alone, the flowers mature in a way that ensures cross pollination. The male parts mature first and offer said pollen. The female parts of the flower are second to mature. They produce no reward for the beetles but are instead believed to mimic the male parts, ensuring that the beetles will spend some time exploring and thus effectively pollinating the flowers.

It is pretty neat to think that you don't necessarily have to track down a dawn redwood or a gingko to see a plant that has survived major extinction events. You can find magnolias very close to home with a keen eye. Looking at one, knowing that this is a piece of biology that has worked for millennia, is quite astounding in my opinion.

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

Meeting Amborella trichopoda

When I found out I would be seeing a living Amborella, a lump formed in my throat. There I was standing in one of the tropical houses at the Atlanta Botanical Garden trying to keep my cool. No amount of patience was ample enough to quell my excitement. How was I going to react? How big were these plants? Would I see flowers? Could I touch them? What were they growing in? My curiosity was through the roof.

Naturally this sort of excitement is reserved for those of us familiar with Amborella trichopoda. This strange shrub is not something that would readily stand out against a backdrop of tropical flora. However, if life history and ecology were to be translated into outward appearances, Amborella would likely be one of the most gaudy plants on this planet. What I was about the lay eyes on is the only member of the sole genus belonging to the family Amborellaceae, which is the sole member of the order Amborellales.

Even more exciting is its position on the angiosperm family tree. As flowering plants go, Amborella is thought to be the oldest alive today. Okay, so maybe this shrub isn't the oldest flowering plant in the world. It is likely that at one time, many millions of years ago, there were more representatives of Amborellaceae growing on this planet. Until we turn up more fossil evidence it is nearly impossible to say. Still, Amborella's place in the story of flowering plant evolution is consistently located at the base.

That is not to say that this shrub is by any means primitive. I think the first thing that shocked me about these plants is just how "normal" they appear. Sans flowers, I didn't see much out of the ordinary about them. They certainly look like they belong on our timeline. Without proper training in plant anatomy and physiology, there is little one could deduce about their evolutionary position. Regardless of my ignorance on plant morphology, there is plenty to look at on Amborella.

For starters, Amborella has tracheids but no vessel elements, making its vascular system more like that of a gymnosperm than an angiosperm. Its small flowers are borne in the axils of the evergreen leaves. It has no petals, only bracts arranged into a spiral of tepals. The female flowers consist of 4 to 8 free carpels and do not produce a style. Male flowers look like nothing more than a spiral cluster of stamens borne on short filaments.

If plant anatomy isn't enough to convince you, then the genetic analyses tell a much more compelling story. DNA sequencing consistently places Amborella at the base of the flowering plant family tree. Again, this is not to say that this shrub is by any means "primitive" but rather its lineage diverged long before what we would readily recognize as a flowering plant evolved. As such, Amborella offers us a window into the early days of flowering plants. By comparing traits present in more derived angiosperms to those of Amborella, researchers are able to better understand how the most dominant group of plants found their place in this world.

Another interesting thing happened when researchers looked at the DNA of Amborella. What they found was more than just Amborella genes. Inside the mitochondrial DNA are an unprecedented amount of foreign DNA from algae, lichens and mosses. In fact, an entire chunk of DNA corresponded to an entire mitochondrial genome of a moss! Researchers now believe that this is a case of extreme horizontal gene transfer between Amborella and its neighbors both growing on and around it. Both in the wild and in cultivation, Amborella is covered in a sort of "biofilm." Whether or not such gene transfer has assisted in the conservatism of this lineage over time remains to be seen.

At this point you may be asking how this lineage has persisted for over 130 million years. For the most part, it is probably due to chance. However, there is one aspect of its ecology that really stands out in this debate and that is its geographic distribution. Amborella is endemic to Grande Terre, the main island of New Caledonia. This is a very special place for biodiversity.

New Caledonia is a small fragment of the once great super-continent Gondwana. New Caledonia, which was part of Australia at that time, broke away from Gondwana when the super-continent began to break up some 200-180 million years ago. New Caledonia then broke away from Australia some 66 million years ago and has not been connected to another land mass since. A warm, stable climate has allowed some of the most unique flora and fauna to persist for all that time. Amborella is but one of the myriad endemic plants that call New Caledonia home. For instance, 43 species of tropical conifers that grow on these small islands are found nowhere else in the world. The whole region is a refugia of a long lost world.

Being a biodiversity hot spot has not spared New Caledonia from the threats of modern man. Mining, agriculture, urbanization, and climate change are all threatening to undo much of what makes this place so unique. The loss of a species like Amborella would be a serious blow to biodiversity, conservation, and the world as whole. We cannot allow this species to exist only in cultivation. New Caledonia is one place we must desperately try to conserve. Meeting this species has left a mark on me. Being able to observe living Amborella up close and personal is something I will never forget as my chances of seeing this species in the wild are quite slim. I am so happy to know that places like the Atlanta Botanical Garden are committed to understanding and conserving this species both in the wild and in cultivation. For now Amborella is here to stay. Long may it be that way.

Further Reading:

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