There's a Pepper Inside My Pepper!

We received an intriguing surprise the other night while prepping dinner. We cut open an unassuming bell pepper (Capsicum annuum) only to find a small, yet perfectly formed pepper inside! It seemed to be attached to the placenta along with the seeds. This was the first time we had ever encountered this, but a quick internet search revealed it isn’t necessarily a rare phenomenon. What was going on with this fruit that caused it to form another fruit within?

The quick answer is parthenocarpy or the formation of fruit without fertilization. Indeed, when we cut into this smaller pepper, there were no seeds inside. Some have taken to calling this phenomenon “internal proliferation,” but the question remained of what caused it to occur in the first place? It can be intentionally induced to produce seedless varieties of various fruits, however, given the “parent” pepper had plenty of seeds, this pepper within a pepper didn’t seem too intentional.

Another internet search revealed that this is an undesirable trait in the pepper trade. Pepper growers will actively select against plants that produce these internal proliferations. However, I have found it difficult to track down any real concrete explanations as to why it happens. Some have suggested that damage to the ovules or other external stressors such as temperature swings can occasionally induce them. Despite the validity of these hypotheses, few have actually bothered to collect and analyze any data.

I did find at least one paper that discussed something called “aberrant ovules” in peppers and their photographs certainly showed internal growths that looked a lot like what we observed in our pepper. These aberrant ovules ranged in appearance from what are essentially mini peppers to mutant blobs of colorful ovary tissue. The only sound conclusions I really took away from their work was that there does seem to be some evidence that genes are involved. They outlined an experiment in which some genetic lineages of peppers were significantly more likely to produce fruits with aberrant ovules than others. That being said, they did not venture much in the way of a trigger for inducing them. That was about as far as I got before I had to attend to other things in life like enjoying the meal we were making.

Regardless of the cause, it was an interesting and unexpected experience to open up this pepper only to find another pepper inside. We ended up eating the little fruit too, and it was just as yummy as the pepper that housed it!

Further Reading: [1] [2]

Let's Talk About Recruitment

Photo by --Tico-- licensed by CC BY-NC-ND 2.0

For any species to be considered successful, it must replace itself generation after generation. We call this process recruitment and it is very important. After all, reproduction is arguably the most fundamental aspect of life in a Darwinian sense. For plants, this can be done either vegetatively or sexually via seeds and spores. Though vegetative reproduction is a fundamental process for many plants around the globe, seed or spore germination is arguably the most important. To truly understand what a plant needs, we have to understand its germination requirements.

Recruitment is a considerable limiting factor for plant populations. In fact, it is the first major bottleneck plants must pass through. It is estimated that a majority of plant mortality occurs during the germination and seedling stages. However, not all plants are equal in this way. Some plants are considered seed or propagule limited whereas others are habitat limited.

If a plant is seed limited, it means that its ability to expand its population or colonize new habitats its limited by the ability of seeds (or spores) to make it to a new location. Once there, nature takes its course and germination occurs with little impediment. If a plant is habitat limited, however, things get a bit more tricky. For habitat limited plants, simply getting seeds to a new location is not enough. Some other aspect of the environment (soil moisture, texture, temperature, disturbance, etc.) limit successful germination. Only when the right conditions are present can habitat limited plants germinate and begin to grow.

Habitat limitation is probably the most common limit to plant establishment. Simply put, not all plants will be successful everywhere. Even the successful growth and persistence of adult plants can be poor predictors of seedling success. Many plants can live for decades or even centuries and the conditions that were present when they germinated may have long since changed. Even the presence of the adults themselves can make a site unsuitable for germination. Think of all of those fire adapted species out there that require the entire community to burn before their seeds will ever germinate.

In reality, it is likely that most plants are habitat limited to some degree. These are not binary categories after all, rather they are aligned along a spectrum of possibilities. The fact that most plants don’t completely take over an area once seeds or spores arrive is proof of the myriad limits to plant establishment. As such, recruitment limitation is extremely important to study. It can make a huge difference in the context of conservation and restoration. Even the successful establishment of adult plants is no guarantee that seedlings stand a chance. Without successful recruitment, all you have left is a nice garden that is doomed to run its course. By understanding the limits to plant recruitment, we can do much more than just improve on our ability to protect and bolster plant populations, we can also gain insights into why so many plants remain rare on the landscape and so few ever rise to dominance.

Photo Credits: [1]

Further Reading: [1] [2]

Pollen Competition

Photo by Martin LaBar licensed under CC BY-NC 2.0

The animal kingdom is rife with sexual conflict. We are all aware of what is going on when two stag deer lock antlers or when a group of male sage grouse flaunt themselves on leks as females look on. But what about plants? Is there sexual conflict among plant species? Whether pollen ends up on a stigma via wind or animal, is there any way for a plant to "choose" who gets to fertilize the ovule?

It turns out, yes, there is. Sexual competition is part of the pollination process. In fact, some of the most familiar floral morphologies may have evolved as a way of weeding out weak paternal lines. To understand this process better, though, we must first quickly review exactly what goes on during pollination.

Photo by Nick Fedele licensed under CC BY-NC-SA 2.0

Pollen is a male gamete. Each grain is haploid and contains only a single copy of a plant’s chromosomes. When a pollen grain lands on a stigma, the grain germinates like a tiny seed, sending down a root-like growth called a pollen tube. This tube grows down into the ovary until it finds an unfertilized ovule. At this point, sperm travels down the pollen tube where it can unite with the ovule, thus forming a seed.

By CNX OpenStax licensed under CC BY 4.0

It’s the formation of this pollen tube that introduces the idea of competition among pollen grains. Again, whether by wind or animal, the pollen arriving to a new plant generally doesn't come from a single individual. Pollen from many potential paternal lines can arrive all at once. As such, the race to fertilize the ovules can be quite intense, and this is where competition begins.

Remember, pollen only contains a single set of chromosomes from the parent plant, thus all alleles, both functioning and deleterious, are represented. During the growth of the pollen tube, upwards of 60% of the pollen genome is actively transcribed. Any pollen containing lots of deleterious alleles is going to have a much harder time competing with pollen grains that have fewer deleterious alleles. Their tubes have a harder time making it to the ovules in time to fertilize them.

Photo by Dartmouth Electron Microscope Facility, Dartmouth College

It is thought that the length of the style (the stem connecting the stigma to the ovaries) may also provide a sort of "proving ground" for pollen too. For instance, picture the flowers of a lily or a mallow. Those long, slender styles may actually be acting like a race track. Only the pollen with the best selection of genetic material will be able to grow their pollen tubes fast enough to reach the ovules, leaving the weaker competition in the dust. In this way, plants may actually be sorting out stronger paternal lines, which makes sense for sessile organisms that can't see.

As with everything in nature, there is far more nuance to this than what I have outlined above. Much work is being done to test some of the earlier assumptions and data surrounding this concept of pollen competition. It certainly happens but the degree to which any given species utilizes such methods is up for debate. Still, it paints a much more interesting picture of mate selection in plants. Static, plants are not!

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

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

The Traveler's Palm

© CEphoto, Uwe Aranas licensed under CC BY-SA 3.0

This nifty looking tree is commonly referred to as the traveler's palm (Ravenala madagascariensis). In reality, it is not a palm at all but rather a close cousin of the bird of paradise plants (Strelitziaceae). It is endemic to Madagascar and the only member of its genus. Even more fascinating is its relationship with another uniquely Madagascan group - the lemurs. But first we must ask, what's in a name?

The name "traveler's palm" has two likely explanations. The first has to do with the orientation of that giant fan of leaves. The tree is said to align its photosynthetic fan in an east-west orientation, which can serve as a crude compass, allowing weary travelers to orient themselves. I found no data to support this. The other possibility comes from the fact that this tree collects a lot of water in its nooks and crannies. Each of its hollow leaf bases can hold upwards of a quart of rain water! Get to it quick, though, because these water stores soon stagnate.

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

Flowers are produced between the axils of the leaves and closely resemble those of its bird of paradise cousins. Closer observation will reveal that they are nonetheless unique. For starters, they are large and contained within stout green bracts. Also, they are considerably less showy than the rest of the family. They don't produce any strong odors but they do fill up with copious amounts of sucrose-rich nectar. Finally, the flowers remain closed, even when mature and are amazingly sturdy structures. It may seem odd for a plant to guard its flowers so tightly until you consider how they are pollinated.

It seems fitting that an endemic plant like the traveler's palm would enter into a pollination syndrome with another group of Madagascar endemics. As it turns out, lemurs seem to be the preferred pollinators of this species. Though black lemurs, white fronted lemurs, and greater dwarf lemurs have been recorded visiting these blooms, it appears that the black-and-white ruffed lemur manages a bulk of the pollination services for this plant.

Watching the lemurs feed, one quickly understands why the flowers are so stout. Lemurs force open the blooms to get at the nectar inside. The long muzzles of the black-and-white ruffed lemur seem especially suited for accessing the energy-rich nectar within. The flowers themselves seem primed for such activity as well. The enclosed anthers are held under great tension. When a lemur pries apart the petals, the anthers spring forward and dust its muzzle with pollen. Using both its hands and feet, the lemur must wedge its face down into the nectar chamber in order to take a sip. In doing so, it inevitably comes into contact with the stigma. Thus, pollination is achieved. Once fertilized, the traveler's palm produces seeds that are covered in beautiful blue arils.

Photo by Jeffdelonge licensed under CC BY-SA 3.0

All in all, this is one unique plant. Though its not the only plant to utilize lemurs as pollinators, it is nonetheless one of the more remarkable examples. Its stunning appearance has made it into something of a horticultural celebrity and one can usually find the traveler's palm growing in larger botanical gardens around the world. Though the traveler's palm itself is not endangered, its lemur pollinators certainly are. As I have said time and again, plants do not operate in a vacuum. To save a species, one must consider the entirety of its habitat. This is why land conservation is so vitally important. Support a land conservancy today!

Photo Credits: [1] [2]

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

Red or White?

Photo by Msact at English Wikipedia licensed under CC BY-SA 3.0

Who doesn't love a nice oak tree? One cannot overstate their importance both ecologically and culturally. Although picking an oak tree out of a lineup is something many of us are capable of doing, identifying oaks to species can be a bit more challenging. This is further complicated by the fact that oaks often hybridize. Still, it is likely you have come across some useful tips and tricks for narrowing down your oak choices. One such trick is distinguishing between the red oaks and the white oaks. If you're anything like me, this is something you took for granted for a while. Is there anything biologically or ecologically meaningful to such a split?

In short, yes. However, a true appreciation of these groups requires a deeper look. To start with, oaks are members of the genus Quercus, which belongs in the family Fagaceae. Globally there are approximately 400 species of oak and each falls into one of three categories - the red oaks (section Lobatae), the white oaks (section Quercus), and the so-called "intermediate" oaks (section Protoblanus). For the sake of this article, I will only be focusing on the red and white groups as that is where most of the oak species reside. The intermediate oak group is made up of 5 species, all of which are native to the southwestern United States and northwestern Mexico.

As is common with oak identification, reliable techniques for distinguishing between the two groups can be tricky. Probably the most reliable feature is located on the inner surface of the acorn cap. In white oaks, it is hairless or nearly so, whereas in red oaks, it is covered in tiny hairs. Another useful acorn feature is the length of time it takes them to germinate. White oak acorns mature in one season and germinate in the fall. As such, they contain lower levels of tannins. Red oak acorns (as well as those of the intermediate group) generally take at least two seasons to mature and therefore germinate the following spring. Because of this, red oak acorns have a much higher tannin content. For more information on why this is the case, read this article.

View fullsize

Tyloses in white oak xylem. Photo by Dr. Michael Rosenthal, Technische Universität Dresden via Rosenthal, M.; Bäucker, E. (2013): Zur Anatomie des Holzes der Weiß-Eichen. Rasterelektronenmikroskopische Bildtafeln zu den drei holzanatomischen Schnittrichtungen. licensed under CC BY-SA 3.0

Less apparent than acorns is the difference in the wood of red and white oaks. The wood of white oaks contains tiny structures in their xylem tissues called tyloses. These are absent from the wood of red oaks. The function of tyloses are quite interesting. During extreme drought or in the case of some sort of infection, they cut off regions of the xylem to stop the spread of an embolism or whatever may be infecting the tree. As such, white oaks tend to be more rot and drought resistant. Fun fact, tyloses are the main reason why white oak is used for making wine and bourbon barrels as it keeps them from leaking their contents.

More apparent to the casual observer, however, is leaf shape. In general, the white oaks produce leaves that have rounded lobes, whereas the red oaks generally exhibit pointed lobes with a tiny bristle on their tips. At this point you may be asking where an unlobed species like shingle oak (Quercus imbricaria) fits in. Look at the tip of its leaf and you will see a small bristle, which means its a member of the red oak group. Similarly, the buds of these two groups often differ in their overall shape. White oak buds tend to be smaller and often have blunted tips whereas the buds of red oaks are generally larger and often pointed.

Tricky leaves of the shingle oak (Quercus imbricaria). Note the bristle tip! Photo by Greg Blick licensed under CC BY-NC-ND 2.0

Despite this broad generalizations, exceptions abound. This is further complicated by the fact that many species will readily hybridize. Quercus is, after all, a massive genus. Regardless, oaks are wonderful species chock full of ecological and cultural value. Still, oak appreciation is something we all need more of in our lives. I encourage you to try some oak identification of your own. Get outside and see if you can use any of these tricks to help you identify some of the oaks in your neighborhood.

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

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

Seed Anchor

[SOURCE]

Epiphytic plants live out their entire lives on the trunks or branches of trees. Using their roots, they attach themselves tightly to the bark. Spend any amount of time in the tropics and it will become quite clear that such a lifestyle has been very successful for a plethora of different plant families. Still, living on a tree isn't easy. Epiphytic plants must overcome harsh conditions among or near the canopy.

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

One of the biggest challenges these plants face starts before they even germinate. This is especially true for orchids. Orchid seeds are more like spores than they are seeds. They are so small that thousands could fit inside of a thimble. Upon ripening, the dust-like seeds waft away on the slightest breeze. In order for epiphytic species to germinate and grow, their seeds must somehow anchor themselves in place on a trunk or branch. Inevitably most seeds are doomed to fail. They simply will not land in a suitable location. It stands to reason then that any adaptation that increases their chances of finding the right kind of habitat will be favored. That's where the strange coils on the tip of Chiloschista seeds, a genus of leafless orchids native to southeast Asia, New Guinea, and Australia, come in. For these orchids, this process is aided by some truly unique seed morphology.

Unlike most orchid seeds that are nothing more than a thin sheath surrounding a tiny embryo, the seeds of Chiloschista have additional parts. These "appendages," which are specialized seed coat cells, are tightly wound into coils. Upon contact with water, these coils shoot out like tiny grappling hooks that grab on to moss and bark alike. In doing so, they anchor the seed in place. By securing their hold on the trunk or branch of a tree, the seeds are much more likely to germinate and grow. This is one of the most extreme examples of seed specialization in the orchid family.

Photo Credit: [1] [2]

Further Reading: [1]

On Crickets and Seed Dispersal

Photo by Vojtěch Zavadil licensed under CC BY-SA 4.0

The world of seed dispersal strategies is fascinating. Since the survival of any plant species requires that its seed find a suitable place to germinate, it is no wonder then that there are myriad ways in which plants disseminate their propagules. Probably my favorite strategies to ponder are those involving diplochory. Diplochory is a fancy way of saying that seed dispersal involves two or more dispersal agents. Probably the most obvious to us are those that utilize fruit. For example, any time a bird eats a fruit and poops out the seeds elsewhere, diplochory has happened.

Less familiar but equally as cool forms of diplochory involve insect vectors. We have discussed myrmecochory (ant dispersal) in the past as well as a unique form of dispersal in which seeds mimic animal dung and are dispersed by dung beetles. But what about other insects? Are there more forms of insect seed dispersal out there? Yes there are. In fact, a 2016 paper offers evidence of a completely overlooked form of insect seed dispersal in the rainforests of Brazil. The seed dispersers in this case are crickets.

Yes, you read that correctly - crickets. Crickets have been largely ignored as potential seed dispersers. Most are omnivores that eat everything from leaves to seeds and even other insects. One report from New Zealand showed that a large species of cricket known as the King weta can disperse viable seeds in its poop after consuming fruits. However, this is largely thought to be incidental. Despite this, few plant folk have ever considered looking at this melodic group of insects... until now. 

The team who published the paper noticed some interesting behavior between crickets and seeds of plants in the family Marantaceae. Plants in this group attach a fleshy structure to their seeds called an aril. The function of this aril is to attract potential seed dispersers. By offering up seeds from various members of the family, the research team were able to demonstrate that seed dispersal by crickets in this region is quite common. Even more astounding, they found that at least six different species of cricket were involved in removing seeds from the study area. What's more, these crickets only ate the aril, leaving the seed behind.

[SOURCE]

The question of whether this constitutes effective seed dispersal remains to be seen. Still, this research suggests some very interesting things regarding crickets as seed dispersal agents. Not only did the crickets in this study remove the same amount of seeds as ants, they also removed larger seeds and took them farther than any ant species. Since only the aril is consumed, such behavior can seriously benefit large-seeded plants. Also, whereas ant seed dispersal occurs largely during daylight hours, cricket dispersal occurs mostly at night, thus adding more resolution to the story of seed dispersal in these habitats. I am very interested to see if this sort of cricket/seed interaction happens elsewhere in the world.

Photo Credits: [1] [2]

Further Reading: [1]

Microclimates

How can we know exactly when and where a seed will germinate? The simple answer is we can't. Much to the chagrin of anyone who has ever tried to grow plants from seed, there seems to be an endless amount of obstacles between deposition of seed and whether or not it will germinate. At the end of the day, it seems that the land will decide where a plant is going to grow. Because of this, the seed to seedling stage of a plants life is the first and greatest bottleneck in the patterns we see in plant communities around the world. 

For some plants its easy. If seed makes it to a location, the plants will grow. For many (and I mean many) plant species, habitat means everything. Anyone interested in growing plants must never overlook microclimates. We all know what climate means. At least we should. Just like entire regions can have their own climates, so too can different parts of the landscape. It could be that snow melt happens slower in one area, making the ground slightly more saturated. Perhaps there is a layer of clay underneath helping to hold water longer. If you are a high desert or alpine species, perhaps a small rock or boulder provides just enough protection from violent winds. It could even be a clump of moss or a shrubs that shelters a seedling long enough for it to establish itself. Heck, it could just as easily be a rusted out old can in the middle of the Mojave Desert.   

The are limitless variations on the theme but they are all too often overlooked. There are few ways of predicting what will work and what won't. On more than one occasion I have forgotten about seeds planted years ago only to have the plants suddenly appear out of nowhere. As with everything in nature, these things are dynamic. What ecology is in need of are more studies that investigate the recruitment limitations of individual or groups of species. We need ecologists speaking with restoration practitioners and vice versa. We need to keep in mind that organisms can inform theory and then some. 

Photo Credit: Zachary Cava (https://www.flickr.com/photos/101789078@N06/)

Further Reading:

http://www.jstor.org/stable/1514074?seq=1#page_scan_tab_contents

Seeds That Plant Themselves

Photo by Matt Lavin licensed under CC BY-SA 2.0

With March (and hopefully spring) just around the corner here in the northern hemisphere, I have been thinking a lot about my garden plans. Winter is a time to get your hands on seeds. What a wonderful thing seeds are. They carry within them the genetic blueprints for building a plant. They are also the means by which most plants get around. Each seed has the potential to start a new generation somewhere else. We are well aware of the myriad ways in which plants equip their seeds for dispersal but the investment doesn't stop there. Many plant species produce seeds that maximize the likelihood of successful germination once dispersed. The ways in which this is done are as diverse as they are interesting.

One of the most remarkable examples of this involves hair-like structures called awns. Awns can be bristly and thus can become tangled in the fur or feathers of an animal. Once on the ground, some awns serve a different purpose. They can be rather sensitive to humidity. This is referred to as "hygroscopic." Hygroscopic awns will begin to twist when humidity levels rise. This movement will actually drill the seed down into the soil where it can safely germinate. Many grasses as well as some geranium seeds behave in this way.

Other seeds have awns or pappuses (hairs) that point backward at an angle which, once driven into the soil, prevent the seed from being pushed back out. This is especially useful when the young roots begin pushing their way down into the soil. Others have pappuses that expand and contract with humidity, placing the seed at a favorable angle for germination when moisture levels are just right. These adaptations are commonly found in species of Leontodon, Taraxacum, Sonchus, Senecio, and Erigeron. Some plants even produce seeds with hairs that become mucilaginous when wet, literally gluing them to the surrounding soil. This adaptation can be seen in Polemonium viscosum.

A seed is an investment for the future. Being static organisms, plants rely on subsequent generations to maintain their presence in and migrate into new habitats. Countless seeds are produced and only a handful will ever survive to flower and repeat the process. Despite these odds, plants are nonetheless incredibly successful.

Photo Credit: Matt Lavin (http://bit.ly/1Bnh4oq)

Further Reading:
http://www.jstor.org/stable/2258879…

http://www.jstor.org/stable/1940966…