Going Veg With Nepenthes ampullaria

Photo by Bernard DUPONT licensed under CC BY-SA 2.0

Carnivory in the plant kingdom is an interesting evolutionary adaptation to living in nutrient poor environments. It has arisen in only a handful of different plant families and indeed, the genera that exhibit it are considered highly derived. There is something to be said about a sessile organism that can take down mobile prey at the rate that most carnivorous plants do.

Perhaps part of our fascination with these botanical wonders stems from their move towards dietary habits not unlike our own. The reason for their predatory behavior is to acquire nutrients like nitrogen and phosphorus. Without these essential nutrients, life as we know it would not exist. It is no wonder then that carnivorous plants have evolved some very interesting ways of getting them into their tissues and to me, there is nothing more peculiar than the way in which Nepenthes ampullaria gets its much needed nitrogen fix.

A rather widespread species, N. ampullaria is at home in the understory of the rain forests of the southeast Asian islands. It differs from its carnivorous cousins in a multitude of ways. For starters, the pitchers of N. ampullaria are oddly shaped. Resembling an urn, they sit in dense clusters all over the jungle floor, below the rest of the plant. Unlike other Nepenthes, the pitchers have only a small, vestigial lid with no nectar glands. Finally, the slippery, waxy surface that normally coats the inside of most Nepenthes pitchers is absent in the pitchers of N. ampullaria. All of these traits are clues to the unique way in which this species has evolved to acquire nitrogen.

N. ampullaria doesn't lure and digest insects. Instead, it relies on leaf litter from the forest canopy above for its nutritional needs. The urn-like shape, lack of a hood, and clustered growth enable the pitchers to accumulate considerable amounts of leaf litter in the pitchers. Because the pitchers are relatively long lived for a Nepenthes, lasting upwards of 6 months, they offer up a nice microhabitat for a multitude of insect and even frog larvae. The collective group of organisms living within the pitchers are referred to as an inquiline community.

Over time, an inquiline community develops in each of the pitchers. This is the key to the success of N. ampullaria. As the inquiline organisms breakdown the leaf litter, they release copious amounts of nitrogen-rich waste. The pitchers can then absorb this waste and begin to utilize it. At least one study found that an individual plant can obtain 35.7% of its foliar nitrogen in this manner. It has also been demonstrated that the pitchers actively manipulate the pumping of hydrogen ions into the fluid within to keep it less acidic than that of other Nepenthes.

I don't know if I would consider this a case of herbivory as the nitrogen is still coming from an animal source but it is nonetheless an interesting adaptation. Instead of using valuable resources on actively digesting its own prey, N. ampullaria is getting other organisms to do the work for it. Not too shabby.

Further Reading:

http://bit.ly/1IRbYG9

http://jxb.oxfordjournals.org/content/61/5/1365

http://link.springer.com/article/10.1007/s004420050390

http://bit.ly/1S10oej

Shady Spines

Tephrocactus articulatus. Photo by Frank Vincentz licensed under CC BY-SA 3.0

Fondling cacti with your bare hands is often ill-advised. These spiny plants are icons of plant defense mechanisms. Cactus spines are actually modified leaves/bud scales. They develop from a bundle of cells called "primordia" that are nearly indistinguishable from leaf primordia. Unlike leaves, however, cactus spines are not made up of living tissue. The genes for leaf development are shut off in these cells and instead, genes for wood fibers are ramped up, creating the stiff structures many of us have had to pry out of our skin.

It is easy to assume that spines are simply there for defense. For a lot species they certainly do the trick. However, for many other species, spines serve another important purpose - they provide shade. This is exemplified by the fact that cacti growing in rainforests and cloudy highlands often have reduced or no spines at all.

For cacti living in the sun-baked regions of the world, sunburn is a serious issue to contend with. Full sunlight can damage sensitive photosynthetic machinery and while intense UV rays wreak havoc on the genome. As such, any adaptation that can shelter these sensitive tissues to some degree is advantageous.

Cephalocereus senilis. Photo by Drew Avery licensed under CC BY 2.0

Spines also buffer the cactus from huge temperature swings. Think of fuzzy or papery spines as a sort of blanket covering the cactus. These spines create a boundary between air immediately surrounding the cactus and the cold nighttime air of these arid climates. This insulation can come in handy as desert temperatures can drop quite low when the sun goes down.

Another benefit spines have is to catch and direct water to the base of the plant. Rain is often scarce in these habitats so when it does occur, a cactus needs to be ready. Water collects on the spines and then runs down to the base. They also act as dew catchers, causing water vapor to condense on their surfaces. In this way, cacti are able to take advantage of every last drop available.

Though they certainly offer some protection, many of these shade spines are too thin and flexible to deter a hungry herbivore. That is where secondary compounds come into play. It is no wonder why some cacti are extremely toxic to herbivores. Whether they are for shade, protection, or water harvesting, cacti spines have managed to capture our imagination and knowing a bit more about their function makes these plants even more impressive.

Photo Credit: [1] [2]

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

Fall Leaves of the Putty-Root Orchid

Whereas most plants here in the Northern Hemisphere have largely geared down for the long winter, there is one species that has only recently begun a new stage of growth. Though it may seem damaging to produce leaves when a hard frost is just around the corner, that is exactly what this plant is doing. What's even more bizarre is that the plant in question is an orchid.

The putty-rood orchid (Aplectrum hyemale) may seem strange to most. Though it flowers during the same time as most of our terrestrial orchids (May through June), its display can be hard to track down. In fact, lacking any knowledge of a specific location, it is more likely that you will stumble across one before you pick it out of the hustle and bustle on the forest floor.

Flowering occurs at a different time than leaf out. The solitary flower stalk gives way to a single leaf starting in late summer or early fall. Why the heck would this plant start its photosynthetic lifecycle when everything else is about ready to go dormant? The answer is competition. Summer is not a bright season for those growing on the forest floor. This is especially true for a plant that only produces a single leaf.

What the putty-root is doing with its oddly timed leaf production is taking advantage of a dormant canopy. With trees and herbaceous leaves out of the way, the putty-root is able to soak up as much sun as it can get. This is a similar strategy adopted by spring ephemerals around the globe. But what does the plant have to gain from having leaves in the fall? Why not wait until spring to leaf out?

As it turns out, it simply doesn't have to. The photosynthetic machinery within the leaves of the putty-root perform exceptionally well at low temperatures. Whereas most plants simply can't photosynthesize when it starts getting too cold, the putty-root is able to photosynthesize at temperatures as low as 2° C (35.6° F)! Not only does this enable the plant to get a jump start come spring, its also able to make food throughout most of fall and even early winter.

There does seem to be a limit to this. Once temperatures drop below 2° C, the machinery can't keep up and photosynthesis grinds to a halt. This is further complicated by the fact that the leaves are often buried under snow for months at a time. Certainly its mycorrhizal associations help feed the plant, even when it isn’t actively photosynthesizing. Regardless, this strategy is a great way of getting an extra kick while everything else is slowing down. Stories such as this bring to mind the story of the tortoise and the hare. Sometimes slow and steady really does win the race!

Photo Credit: Lance Merry (www.lancemerry.com)

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

An Introduction to Cephalotus follicularis - A Strange Carnivore From Australia

Photo by H. Zell licensed under CC BY-SA 3.0

In a small corner of western Australia grows a truly unique carnivorous plant. Commonly referred to as the Albany pitcher plant, Cephalotus follicularis is, evolutionarily speaking, distinct among the pitcher plants. It is entirely unrelated to both the Sarraceniaceae and the Nepenthaceae.

This stunning case of convergent evolution stems from similar ecological limitations. Cephalotus grows in nutrient poor areas and thus must supplement itself with insect prey. It does so by growing modified leaves that are shaped into pitchers. The lid of each pitcher has two main functions. It keeps rain from diluting the digestive enzymes within and it also confuses insects.

A close inspection of the lid will reveal that it is full of clear spots. These spots function as windows, allowing light to penetrate, which confuses insects that have landed on the trap. As they fly upwards into the light, they crash into the lid and fall back down into the trap.

Photo by Lucas Arrrrgh licensed under CC BY-NC-ND 2.0

The relationship of Cephalotus to other plants has been the object of much scrutiny. Though it is different enough to warrant its own family (Cephalotaceae), its position in the greater scheme of plant taxonomy originally had it placed in Saxifragales. Genetic analysis has since moved it out of there and now places it within the order Oxalidales. What is most intriguing to me is that the closest sister lineage to this peculiar little pitcher plant are a group of trees in the family Brunelliaceae. Evolution can be funny like that.

Regardless of its relationship to other plants, Cephalotus follicularis has gained quite a bit of attention over the last few years. Its strange appearance and carnivorous habit have earned it a bit of stardom in the horticultural trade. A single specimen can fetch a hefty price tag. As a result, collecting from wild populations has caused a decline in numbers that are already hurting due to habitat destruction. Luckily they are easy to culture in captivity, which will hopefully take pressure off of them in the wild.

What's more, the loss of Cephalotus from the wild is hurting more than just the plant. A species of flightless, ant-mimicking fly requires Cephalotus pitchers to rear its young. They don't seem to mind growing up in the digestive enzymes of the pitchers and to date, their larvae have been found living nowhere else. If you are lucky enough to grow one of these plants, share the wealth. Captive reared specimens not only take pressure off wild populations, they are also much hardier. Lets keep wild Cephalotus in the wild!

Photo by Holger Hennern licensed under CC BY-SA 3.0

Photo Credits: Holger Hennern (Wikimedia Commons) and Lucas Arrrrgh (https://www.flickr.com/photos/chug/2121092119/)

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

A Fern With Flower Genes - An Odd Case of Horizontal Gene Transfer

Photo by Aaron Carlson licensed under CC BY-SA 2.0

When researchers at Harvard decided to take a look at the genome of the rattlesnake fern (Botrypus virginianum) they found something completely unexpected. Whereas one set of genes they looked at placed this species firmly in the family to which it belongs, Ophioglossaceae, two other genes placed it in the Loranthaceae, a completely unrelated family of flowering plants. What are flowering plant genes doing in a fern?

The rattlesnake fern is a ubiquitous species found throughout the northern hemisphere. It is believed to have evolved in Asia and then radiated outward using ancient land bridges that once connected the continents. At some point before this radiation occurred, the rattlesnake fern picked up some genes that were entirely foreign.

Horizontal gene transfer, the transfer of genes from one organism to another without reproduction, is nothing new in nature. Bacteria do it all the time. Even plants dabble in it every now and then. The surprising thing about this recently documented case is that it is the first discovery of horizontal gene transfer between an angiosperm and a fern. Up until this point, examples within the plant realm have been seen between ferns and hornworts as well as some parasitic plants and their hosts.

This is why the rattlesnake fern genome is so interesting. How did this occur? Though there is no way of telling for sure, researchers believe that one of two things could have happened. The first involves root parasitism. The family Loranthaceae is home to the mistletoes, a group of plants most famous for their parasitic nature. Although the majority of mistletoe species are stem parasites, at least three genera utilize root parasitism. It could be that an ancient species of mistletoe transferred some genes while parasitizing a rattlesnake fern.

This scenario seems to be the least likely of the two as no representatives of the root parasitic mistletoes currently exist in Asia, though it is entirely possible that some did at one time. The other possibility doesn't involve parasitism at all but rather fungi. Rattlesnake ferns are obligate mycotrophs and thus cannot live without certain species of mycorrhizal fungi. Perhaps the transfer of genes was achieved indirectly via a shared mycorrhizal network. This hypothesis is especially tantalizing because if it is found to be true, it would help explain many other examples of horizontal gene transfer that currently lack a mechanism. Only time and more research will tell.

Photo Credit: Aaron Carlson (http://bit.ly/1OAVhNZ)

Further Reading:
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC1560187/

Sticky Friend

Photo by David A. Hofmann licensed under CC BY-NC-ND 2.0

We have all had encounters with sticky plants. Outside of being an interesting sensory experience, the sticky nature of these floral entities would appear to have some evolutionary significance. Considering the cost of producing the glandular trichomes responsible for their stickiness, function is a reasonable question to ask about. For anyone who has taken the time to observe such plants, you will have undoubtedly noticed that insects tend to get stuck to them.

For carnivorous plants, the utility of these glands is readily obvious - trapped insects become food. Even non-carnivores like Roridula gain a nutrient benefit in the form of nutrient-rich feces deposited around the plant by specialized carnivorous bugs that consume trapped insects. However, there are many species of plants out there that fall under the category of "sticky" and a new paper explores this in a more general way.

The serpentine columbine (Aquilegia eximia) is endemic to the Coastal Range of California and it is indeed quite sticky. Its surfaces are covered in glandular hairs. Any given plant can be covered in insects unfortunate enough to come into contact with it. However, it is not a carnivore. As such, researchers wanted to see what benefits, if any, the columbine gained from producing these glands.

By manipulating the amount of insects that were stuck to each plant, researchers found that plants without "victims" actually received more insect damage. The key to this mystery were predators. Plants with lots of trapped victims had more predatory bugs hanging around. These predators, when present, reduced herbivory by deterring other insects that were too large to get stuck. What's more, most of the benefits were observed in the flower buds, which means predators increased the overall reproductive fitness of the serpentine columbine. If the columbine did not trap small insects, these predators would have no reason to hang around.

These predatory bugs were by no means specific to the columbine. In fact, observation of the surrounding plant community found that these predatory insects were present on other sticky genera such as Arctostaphylos, Hemizoni, Holocarpha, Calycidenia, Cordelanthus, Castilleja, Mimulus, Trichostema, and Grindelia. This suggests that the relationship between sticky plants and these generalist predators is more widespread than previously thought. It may also offer a unique window into one possible driver behind the evolution of carnivory in plants.

Photo Credit: David A. Hofmann (http://bit.ly/1l9OtwC)

Further Reading:
http://www.esajournals.org/doi/abs/10.1890/15-0342.1

Orchid Ant Farms

Photo by Scott Zona licensed under CC BY-NC 2.0

I am beginning to think that there is no strategy for survival that is off-limits to the orchid family. Yes, as you may have figured out by now, I am a bit obsessed with these plants. Can you really blame me though? Take for instance Schomburgkia tibicinis (though you may also see it listed under the genera Laelia or more accurately, Myrmecophila). These North, Central, and South American orchids are more commonly referred to as cow-horn orchids because they possess hollow pseudobulbs that have been said to been used by children as toy horns. What is the point of these hollow pseudobulbs?

A paper published back in 1989 in the American Journal of Botany found the answer to that question. As it turns out, ants are quite closely associated with orchids in this genus. They crawl all over the flowers, feeding on nectar. The relationship goes much deeper though. If you were to cut open one of these hollow pseudobulbs, you would find ant colonies living within them. The ants nest inside and often pile up great stores of food and eventually waste within these chambers. The walls of the chambers are lined with a dark tissue that was suspect to researchers.

Using radioactively labeled ants, the researchers found that the orchids were actually taking up nutrients from the ant middens! What's more, nutrients weren't found solely in adjacent tissues but also far away, in the actively growing parts of the roots. These orchids are not only absorbing nutrients from the ants but also translocating it to growing tissues.

While orchids without a resident ant colony seem to do okay, it is believed that orchids with a resident ant colony do ever so slightly better. This makes sense. These orchids grow as epiphytes on trees, a niche that is not high in nutrients. Any additional sources of nutrients these plants can get will undoubtedly aid in their long-term survival. Also, because the ants use the orchids as a food source and a nest site, they are likely defending them from herbivores.

Photo Credit: Scott.Zona (http://bit.ly/1hvWiGX)

Further Reading:
http://www.jstor.org/stable/2444355

Devil's Claws

Proboscidea louisianica

I would like to introduce you to the genus Proboscidea. These lovely, albeit sticky plants are collectively referred to as the Devil's claw plants. The common name comes from the nasty looking seed pods which likely evolved in response to large mammals that once roamed this continent. The genus Proboscidea has traditionally been placed into the sesame family (Pedaliaceae) due to superficial similarities in flower and seed morphology, but more recent work has moved it into the unicorn plant family, Martyniaceae. That's right... unicorn plants.

The entire family is found in the New World, with two species (P. lousianica & P. althaeifolia) hailing from arid parts of the southern portions of North America. At least two other species are readily naturalizing in this region as well. There are some aspects of these species that make them quite interesting to botanists. For starters, the apt name of Devil's claw was bestowed upon this genus because of the bizarre seed pods they produce. Similar to burs, they can become entangled in fur quite readily. The odd thing about this seed dispersal mechanism for some Devil's claws is how big those seed pods are. Until cattle were introduced to this continent, animals large enough to effectively disperse these massive seed pods seemed to be missing, having gone extinct at the end of the last ice age. It is believed that these plants may be an anachronism of this era.

Photo by T.K. Naliaka licensed under CC BY-SA 4.0

Photo by Roger Culos licensed under CC BY-SA 3.0

The flora we are familiar with today spent millennia co-evolving with ice age megafauna like mammoths and giant ground sloths. There is a growing school of thought that many close relationships probably developed over this time and have not yet been lost due to the relatively limited amount of time since the extinction of these large mammals. There are some people who will tell you that the seed pods are "designed" to ensnare small mammals like mice, causing them to die, which then provides the seeds a nutrient-rich, rotting corpses on which to germinate. I have never been able to find any evidence in support of these claims.

Another intriguing anatomical feature of this species are the countless sticky glands that cover the entire plant. These readily ensnare insects that land on or try to climb up the plant. Analysis of the fluids secreted by these glands show evidence of digestive enzymes but the jury still seems to be out on whether or not Devil's claws are undergoing any active carnivorous behavior.

Proboscidea althaeifolia. Public Domain

It is more likely that these glands are a form of defense against insect herbivores and indeed they work quite well. Even a brief run-in with this plant leaves you quite sticky and slimy. It is possible that by ensnaring herbivorous insects, the plant can attract carnivorous insects that will eat the herbivores and then "repay" the devil's claw with nutrient-rich feces. Another possibility is that the glands cause the plant to become covered in sand grains over time. Such sandy armor would get in the way of hungry herbivores. To ad insult to injury, the plant kind of smells. It has been likened to old gym clothes.

These are neat plants. I have had fun growing them in the past. They are an annual but may reseed if care is not taken to removing the seed pods before they pop open. Because of their lively appearance and the unique look of their seed pods, these plants are often grown as horticultural oddities. Be careful though, as they have escaped cultivation outside of their native range and can be considered a noxious weed!

Photo Credit: [1] [2] [3]

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

Aquatic Angiosperm: A Cretaceous Origin?

Via Bernard Gomeza, Véronique Daviero-Gomeza, Clément Coiffardb, Carles Martín-Closasc, David L. Dilcherd, and O. Sanisidro [SOURCE]

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

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

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

Via Bernard Gomeza, Véronique Daviero-Gomeza, Clément Coiffardb, Carles Martín-Closasc, David L. Dilcherd, and O. Sanisidro [SOURCE]

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

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

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

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

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

When a Mutualism Becomes Obligate

Photo by Tony Rodd licensed under CC BY-NC-SA 2.0

Mutualism. The word invokes this warm and fuzzy "you scratch my back and I'll scratch yours" feeling. It is easy to grasp how a mutualism would develop and be maintained. But, in any system, there are bound to be cheaters. Cheaters reduce the fitness of one of the partners so to avoid such things, some species up the ante by resorting to some interestingly "sinister" methods.

Acacias and ants have quite the relationship. Acacias protect themselves by offering ants hollow spines and branches where their colonies can live. They even sweeten the deal via extrafloral nectaries. These are glands on the stems that secrete nectar that the ants eat. In some ant species, this is their only source of food. Needless to say, the ants become highly protective of their acacia trees. They readily attack herbivores and even go as far as to prune away vegetation that may interfere with their host. This seems like a pretty straight forward mutualistic relationship, right?

Ah, but it goes deeper. To make sure that the ants will solely rely on the acacia and are thus completely tied up in the well being of their host, the acacia alters the ants phenotype at birth. Normally these ants have no issues digesting sucrose. Researchers found that the nectar in the extrafloral nectaries contains a protein called "chitinase." Chitinase inhibits the ability of the ants to digest sucrose. When ant eggs hatch into larvae, their first meal is nectar from the extra floral nectaries. Once the larvae ingest this protein they are no longer able to feed on anything other than their hosts nectar. Thus their very survival is completely tied to the Acacia.

I am positive that more examples of such obligate mutualisms abound in nature. We only have to ask the right questions to discover them. It is also interesting considering what we are finding out about our own behavior and how it relates to the microbiome living on and within us. What about human behavior could be described in the context of a relationship similar to ants and acacias?

Photo Credit: Tony Rodd

Further Reading:
http://www.ncbi.nlm.nih.gov/pubmed/24188323

American Persimmon

Photo by Doug McAbee licensed under CC BY-NC 2.0

I will never forget the time I went to the grocery store and bought what I thought were strange tomato varieties. I got home and dug into them only to discover they were not tomatoes at all. I quickly realized the error in my judgment. Instead of the unmistakable flavor of a tomato, what I experienced was something slightly sweet and kind of astringent. I had inadvertently purchased a couple persimmon fruits. I was young and naive so I will cut myself some slack, however, like any good mistake, I was rewarded by the inadvertent introduction to a fascinating fruit I had never experienced before. 

Thinking this to be some strange tropical species, I was surprised to learn that North America does indeed have its own species of persimmon. Known scientifically as Diospyros virginiana, the American persimmon is native to much of the eastern U.S. but is absent north of Pennsylvania. We are lucky, biogeographically speaking, to have this species as the family to which it belongs, Ebenaceae, is predominantly tropical. It is an early successional tree species, often growing on recently abandoned farmland. In the spring this shrubby tree produces small yet attractive white and yellow flowers. American persimmon are dioecious meaning individual trees are either male or female. Their main pollinators are bees.  

As is often seen with many fruiting tree species,  there is a lot of variety between the fruits of different persimmons. They can range in size from small crabapples to the tomato-like fruits we find in the grocery store. There are those who suspect the fruits of the American persimmon to be a throwback to a time when animals like woolly mammoths and ground sloths roamed this continent, dispersing persimmon seeds as they roamed across the terrain. Indeed, fossils of American persimmon have been found in Miocene deposits in areas of Greenland and Alaska which suggests that this species has undergone range contraction, potentially due to the loss of these large seed dispersers. However, modern day evidence would seem to suggest otherwise. Today, much smaller animals like raccoons and opossum seem to do just as good of a job as a larger animal would. It is likely that the constricted range of the American persimmon has more to do with climate than seed dispersal. 

If you have never tried a persimmon before then seek one out and give it a go. If you find them in a grocery store, there is a good chance the fruit belongs to the Asian species (Diospyros kaki). The key to enjoying an American persimmon is making sure its ripe. If you are too early you are going experience some of the worst tannin dry mouth (I honestly don't think I will ever convince my mother to eat another strange fruit again). Either way, this neat species often goes overlooked until it is in fruit. Keep your eye out for fruiting persimmon in your area and report back if you decide to sample some. 

Photo Credit: Doug McAbee (http://bit.ly/1xznvPx)

Further Reading:
http://www.na.fs.fed.us/pubs/silvics_manual/volume_2/diospyros/virginiana.htm

Is it a Fungus? Is it a Forb? No, it's a Tree!

Botanical gardens are winter sanctuaries for a northerner like myself. Winter tree ID can only do so much for me during these times. As such, I try my best to make regular trips to tropical houses wherever and whenever I can. On a recent excursion to the Missouri Botanical Garden, I came across something completely unexpected.

I was perusing their tropical house aptly named "The Climatron." As I rounded a corner I happened to look down and saw what looked like something only a member of the birthwort family (Aristolochiaceae) could produce. There, lying near the ground were a cluster of some of the coolest flowers I have personally laid eyes on.

Photo by Cymothoa exigua licensed under CC BY-SA 3.0

I began searching for the plant that produced them. Up until this point, I have only encountered members of this family in the form of low-lying understory herbs and scrambling vines dangling from the canopy. There were no apparent leaves associated with these flowers and the part of my brain responsible for search images became confused. I traced the flower stems to their place of origin and realized they were attached to the nearest trunk. I followed the trunk upwards and realized that what I had found was in fact a small tree!

The species I was looking at was none other than Aristolochia arborea, a small tree native to the tropical forests of Central America. Needless to say I was floored. There is something to be said about any plant family than can vary this much in size and habit. The coolest aspect about this tree is that, similar to the more herbaceous members of this family, the flowers are produced close to or directly on the forest floor.

A closer inspection of these strange blooms reveals an interesting morphology. It would appear that they are mimicking fungi in the genus Marasimus. Now this could simply be a manifestation of apophenia. Was I seeing patterns where there are none? Of course, this was a job for scientific literature.

It seems I may have been on to something. Botanists agree that in the wild this plant is pollinated by fungus gnats and flies. However, no direct observations of this have ever been made. That being said, the flowers do emit a rather musty smell that could very well be described as "fungal." Regardless, this is an excellent choice of tree to showcase in a botanical garden because stumbling into it like I did led me down an curious path of discovery.

Tree photo credit: Cymothoa exigua (Wikimedia Commons)

Further Reading: [1] [2]

Osage Orange

As a kid I used to get a kick out of a couple trees without ever giving any thought towards what it was. My friend's neighbor had a some Osage orange trees (Maclura pomifera) growing at the end of his driveway. Their houses were situated atop a large hill and the road was pretty much a straight drop down into a small river valley. After school on fall afternoons, we would hang out in my friends front yard and watch as the large "hedge apples" would fall from the tree, bounce off the hood of his neighbor's car (why he insisted on parking there is beyond me) and go rolling down the hill. I never would have guessed that almost two decades later the Osage orange would bring intrigue into my life yet again. This time, however, it would be because of the evolutionary conundrum it presents to those interested in a paleontological mystery...

The fruit of this tree are strange. They are about the size of a softball, they are green and wrinkly, and their insides are filled with small seeds encased in a rather fibrous pulp that oozes with slightly toxic white sap. No wild animal alive today regularly nibbles on these fruits besides the occasional squirrel and certainly none can swallow one whole. Why then would the tree go through so much energy to produce them when all they do anymore is fall off and rot on the ground? The answer lies in the recently extinct Pleistocene megafauna. 

The tree is named after the Osage tribe who used to travel great distances to the only known natural range of this tree in order to gather wood from it for making arrows. It only grew in a small range within the Red River region of Texas. When settlers made it to this continent, they too utilized this tree for things like hedgerows and natural fences. 

What is even stranger is that recent fossil evidence shows that Maclura once had a much greater distribution. Fossils have been found all the way up into Ontario, Canada. In fact, it is believed that there were once 7 different species of Maclura. It was quickly realized that this tree did quite well far outside of its current natural range. Why then was it so limited in distribution? Without the Pleistocene megafauna to distribute seeds, the tree had to rely on flood events to carry the large fruit any great distance. With a little luck, a few seeds would be able to germinate out of the rotting pulp. Botanists agree that the Red River region was a the last stronghold for this once wide ranging species until modern man came on the scene. 

Another clue comes from the toxicity of the fruits. Small animals cannot eat much of it without being poisoned. This makes sense if you are a Maclura relying on large animals as dispersers. You would want to arm your fruit just enough to discourage little, inefficient fruit thieves from making a wasteful meal out of your reproductive effort. However, by limiting the amount of toxins produced in the fruit, Maclura was still able to rely on large bodied animals that can eat a lot more fruit without getting poisoned. Today, with the introduction of domesticated megafauna such as horses and cows, we can once again observe how well these fruits perform in the presence of large mammals. 

Finally, for anyone familiar with Maclura, you will notice that the tree is armed with large spines. Why the heck does a large tree need to arm itself so extravagantly all the way to the top? Again, if you need things like mammoths or giant ground sloths to disperse your seeds, you may want to take some extra precautions to make sure they aren't snacking on you as well. It takes energy to produce spines so it is reasonable to assume that the tree would not go through so much trouble to protect even its crown if there once wasn't animals large enough to reach that high. The Pleistocene megafauna went extinct in what is evolutionarily speaking only the blink of an eye. Trees like the Osage orange have not had time to adapt accordingly. As such, without the helping hand of humans, this tree would still be hanging on to a mere fraction of its former range down in the Red River region of Texas.

Further Reading:
http://plants.usda.gov/core/profile?symbol=MAPO

http://www.americanforests.org/magazine/article/trees-that-miss-the-mammoths/

http://www.plosone.org/article/info:doi%2F10.1371%2Fjournal.pone.0001745

Fiery Peppers - Evolution of the Burn

Photo by Ryan Bushby licensed under CC BY 2.5

Love them or hate them, one must respect the fiery chili pepper. If you're like me then the addition of these spicy fruits can greatly enhance the culinary experience. For others, spice can be a nightmare. Peppers are so commonplace throughout many cultures of the world that it is easy to overlook them. As a plant fanatic, even the simple act of cooking dinner opens the door to so many interesting questions. What is a pepper? Where do they come from? And why are some so spicy?

Peppers evolved in the Americas. The genus to which they belong, Capsicum, is comprised of somewhere around 27 species. Of these, five have been domesticated. They have no relation whatsoever to black pepper (Piper nigrum). Instead, the chili peppers are relatives of tomatoes, potatoes, and eggplants - family Solanaceae.

The fruit that they produce is actually a type of berry. In the wild, Capsicum fruits are much smaller than the ones we buy at the farmers market or grocery store. Centuries of domestication has created such gaudy monsters. The spicy effect one experiences when biting into a pepper is the result of a chemical called capsaicin. It is mainly produced in the placental tissues and the internal membranes. It is in its highest concentrations in the white pith that surrounds the seeds.

As with any fruit, the main goal is seed dispersal. Why then would the plant arm its fruits with fiery capsaicin? The answer to this riddle lies in their wild relatives. As mentioned, the fruits of wild peppers are much smaller in nature. When ripe, they turn bright shades of reds, yellows, and oranges. Their small size and bright coloration are vivid sign posts for their main seed dispersersal agents - birds.

As it turns out, birds are not sensitive to capsaicin. Mammals and insects are, however, and that is a fact not lost on the plants. Capsaicin is there to deter such critters from feeding on the fruits and wasting hard earned reproductive efforts. As such, the well defended fruits can sit on the plant until they are ripe enough for birds to take them away, spreading seeds via their nutrient rich droppings.

It may be obvious at this point that the mammal-deterring properties of Capsicum have been no use on humans. Many of us enjoy a dash of spice in our meals and some people even see it as a challenge. We have bred peppers that are walking a thin line between spicy and dangerous. All of this has been done to the benefit of the five domesticated species, which today enjoy a nearly global distribution. Take this as some food for thought the next time you are prepping a spicy meal.

Photo Credits: Ryan Bushby, André Karwath, and Eric Hunt - Wikimedia Commons

Further Reading:
http://link.springer.com/article/10.1007%2FBF00994601

http://www.jstor.org/stable/4163197…

http://www.press.uchicago.edu/ucp/journals/journal/ijps.html

Mighty Mighty Squash Bees

Photo by MJI Photos (Mary J. I.) licensed under CC BY-NC-ND 2.0

It's decorative gourd season, ladies and gentlemen. If you are anything like me then you should be reveling in the tastes, smells, and overall pleasing aesthetics of the fruit of the family Cucurbitaceae. If so, then you must pay your respects to a hard working bee that is responsible for the sexual efforts of these vining plants. I'm not talking about the honeybee, no no. I am talking about the squash bees. 

If we're being technical, the squash bees are comprised of two genera, Peponapis and Xenoglossa. They are not the hive forming bees we generally think of. Instead, these bees are solitary in nature. After mating (which usually occurs inside squash flowers) the females will dig a tunnel into the ground. Inside that tunnel she places balls of squash pollen upon which she will lay an egg. The larvae consume the protein-rich pollen as they develop. 

The story of squash bees and Cucurbitaceae is a North American story. Long before squash was domesticated, these bees were busy pollinating their wild relatives. As a result, this bee/plant relationship is quite strong. Female squash bees absolutely rely on squash flowers for the pollen and nectar needs of their offspring. In fact, they often dig their brood tunnels directly beneath the plants. 

Because of this long standing evolutionary relationship, squash bees are the best pollinators of this plant family. The flowers open in the morning just as the squash bees are at their most active. Also, because they are so specific to squash, the squash bees ensure that pollen from one squash flower will make it to another squash flower instead of an unrelated plant species. Honeybees can't hold a candle to these native bees. What's more, crowds of eager honeybees may even chase off the solitary squash bees. For these reasons, it is often recommended that squash farmers forgo purchasing honeybee hives for their crops. If left up to nature, the squash bees will do what they are evolutionarily made to do. 

Photo Credit: MJI Photos (https://www.flickr.com/photos/capturingwonder/4962652272/)

Further Reading:
http://www.researchgate.net/profile/Victor_Parra-Tabla2/publication/226134213_Importance_of_Conserving_Alternative_Pollinators_Assessing_the_Pollination_Efficiency_of_the_Squash_Bee_Peponapis_limitaris_in_Cucurbita_moschata_(Cucurbitaceae)/links/549471010cf20f487d2a95b8.pdf

http://www.jstor.org/stable/25084168?seq=1#page_scan_tab_contents

http://extension.psu.edu/plants/sustainable/news/2011/jan-2011/1-squash-bees

Why We See Color

Photo by Francisco Anzola licensed under CC BY 2.0

Seeing the world in trichromatic color is a wonderful thing. I truly feel for those who can't. Humans, by and large, have pretty decent color vision. We have three different kinds of opsins on our cones which allows us to see the variety in hues that we do. It is a trait we share with apes and most Old World monkeys. Why do we possess such a wonderful adaptation? As it turns out, plants were likely the driving factor.

Whereas most mammals tend to have only two different kinds of opsins (dichromacy), the primate lineage from which we evolved developed trichromacy at some point in the past. Why did this happen? The answer may lie in the diet of our common ancestors. As climates changed over time, the common ancestors of Old World monkeys, apes and humans had to constantly adapt to new food sources. A majority of primate diets consist of fruits and leaves. Being able to distinguish between ripe and unripe fruit would be a valuable advantage to have. For our ancestors, dichromacy would have made this quite difficult. Thus the evolution of trichromacy would have incurred quite a selective advantage to our ancestors.

The advantage doesn't end with ripe vs. unripe either. Trichromacy would have also made finding colorful fruits against a backdrop of green much easier as well. Even for the majority of primates that eat leaves, color vision would have been quite useful. Leaves can vary in edibility and even toxicity with age. Being able to tell younger from older leaves could easily make the difference between life and death for these primates. Leaf color is often the only way this can be done. Again, selection for color vision would have quickly spread through these populations. So, the next time you stop to admire a flower or any of the wonderful colors of the world around you, take a moment to think about the fact that plants just might be the reason you can enjoy that wonderful sense.


Further Reading:
http://anthro.palomar.edu/primate/color.htm

http://rspb.royalsocietypublishing.org/content/263/1370/593

http://www.sciencedirect.com/science/article/pii/S0047248402001677

The Deciduous Conifer Conundrum

Broad leaf trees get all the glory come fall. Their dazzling colors put on a display for a few weeks every year that is unrivaled. However, it isn't just broad leaf trees that are preparing for winter in this manner. There are some conifers doing the same. The handful that have evolved this deciduous strategy are just as dazzling as their broad leaf neighbors.

The most famous of these are the larches (genus Larix), however, there are others such as baldcypress (genus Taxodium) and the dawn redwoods (genus Metasequoia). So, why have these conifers evolved to be deciduous? There are likely many reasons these genera utilize this strategy but it most likely comes down to cost versus benefit. Needles that last for years are costly to make despite their advantages. They are no guarantee of success either, especially for the larches, which often grow in areas that experience some of the harshest winters on the planet. Heavy snow pack and deep winter chills can take their toll on conifers and many evergreen species show signs of frost damage and broken limbs from snow loads. The habitats in which deciduous conifers are found can be tough places to eek out a living.

By shedding their needles, the larches can get around these issues a bit. They also tend to grow in swampy areas where getting the nutrients needed for survival can be extra difficult. By producing relatively weak needles that are easily replaced from year to year, trees like larches and cypress may get around having to waste resources on more robust needles. Finally, it should be noted that this strategy is by no means less efficient. These genera do quite fine with their deciduous nature. For the most part, these trees are nonetheless successful and can live for centuries. It is mysteries like these that keep the wonderful world of botany interesting.

Further Reading:
http://www.metla.fi/silvafennica/full/sf36/sf363703.pdf

Blue

Blue is a strange color. This may seem like an odd statement yet, when you think about it, so few things in nature are truly blue. It is estimated that, of all the colors plants utilize to attract pollinators, blue occurs in less than 10% of species. This isn't a pattern restricted to plants either. Blue is an infrequent occurrence throughout the biological world.

When it does appear, the color blue is usually the result of structure rather than pigment. The feathers of a bluejay, the wings of a morpho butterfly, and the sheen of a beetles elytra - these blues owe their vibrancy to refracted light, not pigment. Without light, the crystalline cells responsible for the blue hue would appear dull brown. As light enters their structure, it is bent in a way that gives off blue wavelengths.

The metalic blue hue of these Pollia condensata are the result of refracted light, not pigment. Photo by Juliano Costa licensed under CC BY-SA 3.0

Plants have adopted this strategy as well. The berries of Pollia condensata use a similar crystalline structure that results in blue. However, there are true blue flowers out there. How have species with blue flowers managed to overcome the rarity of blue pigments?

The simple answer is that they haven't. There are no blue pigments in the floral world. Instead, plants utilize what can only be described as an evolutionary hack. Blue flowers obtain their color by doing something we all did in art class, blending pigments (similar to the one true black flower). By producing varying amounts of anthocyanins (the pigments responsible for reds) floral cells are able to make blue flowers.

The anthocyanins can also be tweaked to appear blue. One way of doing this is through changes in pH. The famous blue poppies (Meconopsis grandis), for example, have a defect in the proton pumps found inside their flower cells. This causes the cells to become more basic than acidic, which manifests in blue, rather than purple, flowers. Blue petunias do this as well.

Despite the lack of blue in the floral world, it nonetheless seems to work well when it comes to pollinators. I watched multiple different species of bee visit the flowers of this downy gentian (Gentiana puberulenta). Hummingbirds often visit the amazing floral display produced by the great blue lobelias (Lobelia siphilitica) in my garden. Anyone that has looked over a patch of blue lupine or delphiniums can attest to the success of this color.

Photo Credits: [1]

Further Reading: [1] [2]

CAM Photosynthesis

I was in a lecture the other day and I heard something that made the plant nut inside of me chuckle. The professor was trying to make the point that C3 photosynthesis is the most common photosynthetic pathway on the planet. To do this he said "it is the vanilla pathway." In this context, he was using vanilla as an adjective meaning "plain or ordinary." Of course, this was all very facetious, however, I thought it interesting and funny how, if taken literally, that statement was just plain wrong. 

I have written before about the reproductive ecology of Vanilla orchids (http://bit.ly/1LcC857). They are anything but vanilla the adjective. The other part of the statement that was wrong (again, if taken literally) is that C3 is the photosynthetic pathway of the vanilla orchid. In reality, vanillas are CAM photosynthesizers.

Last week I wrote about the C4 pathway and how it has helped plants in hot, dry places, but the CAM pathway is yet another adaptation to such climates. The interesting thing about CAM photosynthesis is that it separates out the different reactions in the photosynthetic pathway on a temporal basis. 

CAM is short for Crassulacean acid metabolism. It was first described in succulents in the family Crassulaceae. Hence the name. Similar to the C4 pathway, CO2 is taken into the leaves of the plant and stored as an organic acid. This is where the process differs. For starters, having acid hanging around inside your leaves is not necessarily a good thing. CAM plants deal with this by storing it in large vacuoles. That is one reason for the succulent appearance of many CAM species. 

Because these plants so often grow in hot, dry climates, they need to minimize water loss. Water evaporates from holes in the leaves called stomata so to avoid this, these holes must be closed. However, closing the stomata means not letting in any CO2 either. Whereas C4 plants get around this by only opening their stomata during the cooler hours of the day, CAM plants forgo opening their stomata entirely when the sun is up. 

Instead, CAM plants open their stomata at night when the vapor pressure is minimal. This ensures that water loss is also minimal.  Like camels storing water for lean times, CAM plants store CO2 as organic acid to use when the sun rises the next day. In this way, CAM plants can close their stomata all the while the hot sun is baking the surrounding landscape yet still undergo ample photosynthesis for survival. 

Not all orchids do this. In fact, some can switch photosynthetic pathways in different tissues. However, there are many other CAM plants out there including some very familiar species like pineapples, cycads, peperomias, and cacti. If you're like me and prone to talking to your plants, it is probably best to talk to your CAM plants after the sun has set. Not only does it confuse neighbors and friends, it provides them with CO2 when they are actively absorbing it. 

Further Viewing: https://www.khanacademy.org/science/biology/cellular-molecular-biology/photosynthesis/v/cam-plants

There's Water In Them There Rocks!

Photo by José María Escolano licensed under CC BY-NC-SA 2.0

Plants go to great lengths to obtain the necessities of survival. Nowhere is this more apparent than in the desert regions around the world. Amazingly, myriads of plants have adapted to the harsh conditions that deserts offer up. Needless to say, water is a major limiting resource in these climates and many of the adaptations we see in desert plant species have to do with obtaining and holding on to as much water as possible. Some species get around the issue by going dormant whereas others stick it out using deep taproots that plug into the groundwater. A select few others hit the rocks.

Rocks? Well, gypsum to be precise. This interesting mineral is quite common in arid regions throughout the world. What is more interesting is that 20.8% of a gypsum crystal is water. Because of this, it has been suspected that gypsum in the soil could be a potential source of water for plants growing in these regions and a team of researchers out of Spain may have found just that.

Meet Helianthemum squamatum. This distant relative of hibiscus grows throughout the gypsum hills of the Mediterranean region. Unlike other desert plants, it is shallowly rooted. Unlike other shallowly rooted species, H. squamatum doesn't go dormant during the dry summer months. The physiology of this species in the context of the dry environments that it grows offers up quite a conundrum. How does this plant get the water it needs to grow through the hottest, driest months of the year?

By analyzing the isotopic composition of the water within the plant and comparing it to background sources, the team found that 90% of the plants water intake during the dry summer months comes from the crystallization water in gypsum! How is this possible? How does a plant get water from a mineral?

The actual physiological processes involved are not yet understood but there are some running hypotheses. The first has to do with temperature. When gypsum is exposed to temperatures above 40 degrees C, water can be released from the crystalline matrix. It would then be available to the plants via passive uptake. 40 degrees C is not unheard of in these environments. Any water that isn't taken up by the plants could be reincorporated back into gypsum when things cool down at night. Another possibility is that H. squamatum grows its roots into and around the gypsum. Using root exudates, it is possible that the plant is able to dissolve gypsum to some degree, thus unleashing the water within. This may rely on the microbial community associated with the roots. Until further research can be done on this, the jury is still out.

The most exciting aspect of this research is the doors it has now opened in our search for extraterrestrial life. Life as we know it depends on water. Our search for this molecule has us looking for planets in a sweet spot where water can be found in a liquid state. Knowing now that at least some life on our planet is able to obtain water from gypsum broadens the kinds of places we can look. Mars is chock full of gypsum. Just sayin'.

Photo Credit: José María Escolano (http://bit.ly/ZeSVzB)

Further Reading:

http://www.nature.com/ncomms/2014/140818/ncomms5660/full/ncomms5660.html