The Giant Genomes of Geophytes

Canopy plant (Paris japonica) Photo by Radek Szuban licensed under CC BY-NC 2.0

A geophyte is any plant with a short, seasonal lifestyle and some form of underground storage organ ( bulb, tuber, thick rhizome, etc.). Plants hailing from a variety of families fall into this category. However, they share more than just a similar life history. A disproportionate amount of geophytic plants also possess massive genomes. 

As we have discussed in previous posts, life isn't easy for geophytes. Cold temperatures, a short growing season, and plenty of hungry herbivores represent countless hurdles that must be overcome. That is why many geophytes opt for rapid growth as soon as conditions are right. However, they don't do this via rapid cell division. 

Dutchman's breeches (Dicentra cucullaria) emerging with preformed buds.

Instead, geophytes spend the "dormant" months pre-growing all of their organs. What's more, the cells that make up their leaves and flowers are generally much larger than cells found in non-geophytes. This is where that large genome comes into plant. If they had to wait until the first few weeks of spring to start their development, a large genome would only get in the way. Their dormant season growth means that these plants don't have to worry about streamlining the process of cellular division. They can take their time. 

As such, an accumulation of genetic material isn't detrimental. Instead, it may actually be quite beneficial for geophytes. Associated with large genomes are things like larger stomata, which helps these plants better regulate their water needs. The large genomes may very well be the reason that many geophytic plants are so good at taking advantage of such ephemeral growing conditions. 

When the right conditions present themselves, geophytes don't waste time. Pre-formed organs like leaves and flowers simply have to fill with water instead of having to wait for tissues to divide and differentiate. Water is plentiful during the spring so geophytes can rely on turgor pressure within their large cells for stability rather than investing in thick cell walls. That is why so many spring blooming plants feel so fleshy to the touch. 

Taken together, we can see how large genomes and a unique growth strategy have allowed these plants to exploit seasonally available habitats. It is worth noting, however, that this is far from the complete picture. With such a wide variety of plant species adopting a geophytic lifestyle, we still have a lot to learn about the secret lives of these plants.

Photo Credits: [1] [2]

Further Reading: [1]

Plant Architecture and Its Evolutionary Implications

View fullsize

[SOURCE]

I make it a point that during my field season I enjoy my breakfast out on the deck. It is situated about halfway up the canopy of the surrounding forest and offers a unique perspective that is hard to come by elsewhere. Instead of looking up at the trees, I am situated in a way that allows for a better understanding of the overall structure of the forest. Its this perspective that generates a lot of different questions about what it takes to survive in a forested ecosystem, especially as it relates to sessile organisms like plants.

Quite possibly my favorite plants to observe from the deck are the pagoda dogwoods (Cornus alternifolia). Hinted at by its common name, this wonderful small tree takes on a pagoda-like growth form with its stacked, horizontal branching pattern. It is unmistakable against the backdrop of other small trees and shrubs in the mid canopy. The fact that it, as well as many other plant species, can be readily recognized and identified on shape alone will not be lost on most plant enthusiasts.

The fact that diagrams like these exist in tree guides is proof of the utility of this concept.

Even without the proper vocabulary to describe their forms, anyone with a keen search image understands there is a gestalt to most species and that there is more to this than simply fodder for dichotomous keys. The overall form of plants has garnered attention from a variety of disciplines. Such investigations involve fields of study like theoretical and quantitative biology to engineering and biomechanics. It has even been used to understand how life may evolve on other planets. It is a fascinating field of investigation and one worth spending time in the literature. 

Some of the pioneering work on this subject started with two European botanists: Dr. Francis Hallé and Dr. Roelof Oldeman. Together they worked on conceptual models of tree architecture. Using a plethora of empirical studies on whether a tree branches or doesn't, where branches occurs, how shoots extend, how branches differentiate, and whether reproductive structures are terminal or lateral, they were able to reduce the total number of tree forms down to 23 basic architectural models (pictured above). Each model describes the overall pattern with which plants grow, branch, and produce reproductive structures. At the core of these models is the concept of reiteration or the repitition of form in repeatable sub-units. The models themselves were given neutral names that reflect the botanists that provided the groundwork necessary to understand them.  

Despite the fact that these models are based on investigations of tropical tree species, they are largely applicable to all plant types whether they are woody or herbaceous and whether they occur in the temperate zone or the tropics. The models themselves do not represent precise categories but rather points on a spectrum of architectural possibilities. Some plants may be intermediate between two forms or share features of more than one model. It should also be noted that most trees conform to a specific model for only a limited time period during their early years of development. Random or stochastic events throughout a trees life greatly influence its overall structure as it continues to grow. The authors are careful to point out that a trees crown is the result of all the deterministic, opportunistic, and chance events in its lifetime.  

Despite these exceptions, the adherence of most plants to these 23 basic models is quite astounding. Although many of the 23 models are only found in the tropics (likely an artifact of the higher number of species in the tropics than in the temperate zones), they provide accurate reference points for further study. For instance, the restriction of some growth forms to the tropics raises intriguing questions. What is it about tropical habitats that restricts models such as Nozeran's (represented by chocolate - Theobroma cacao) and Aubréville's (represented by the sea almond - Terminalia catappa) to these tropical environments? It likely has to do with the way in which lateral buds develop. In these models, buds develop without a dormancy stage, a characteristic that is not possible in the seasonal climates of the temperate zones. 

View fullsize

Reiteration is an important process in plant architectural development in which plants repeat their basic model. This is especially important in repairing damage. [SOURCE]

Another interesting finding borne from these models is that there doesn't seem to be strong correlations between architecture and phylogeny. Although species within a specific genus often share similar architecture, there are plenty of exceptions. What's more, the same form can occur in unrelated species. For instance, Aubréville's model occurs in at least 19 different families. Similarly, the family Icacinaceae, which contains somewhere between 300 and 400 species, exhibits at least 7 of the different models. Alternatively, some families are architecturally quite simple. For instance the gymnosperms are considered architecturally poor, exhibiting only 4 of the different models. Even large families of flowering plants can be architecturally simplistic. Take the Fabaceae, which is largely comprised of plants exhibiting Troll's model. 

So, at this point the question of what is governing these models becomes apparent. If most plants can be reduced to these growth forms at some point in their life then there must be some aspect of the physical world that has shaped their evolution through time. Additionally, how does plant architecture at the physical level scale up to the level of a forest? Questions such as this are fundamental to our understanding of not only plants as organisms, but the role they play in shaping the world around us. 

Although many scientists have attempted to tackle these sorts of questions, I want to highlight the work of one individual in particular - Dr. Karl Niklas. His work utilizes mathematics to explain plant growth and form in relation to four basic physical constraints:

1) Plants have to capture sunlight and avoid shading their own leaves.

2) Plants have to support themselves structurally.

3) Plants have to conduct water to their various tissues.

4) Plants must be able to reproduce effectively.

Using these basic constraints, Dr. Niklas built a mathematical simulation of plant evolution. His model starts out as a "universe" containing billions of possible plant architectures. The model then assesses each of these forms on how well they are able to grow, survive, and reproduce through time. The model is then allowed to change environmental conditions to assess how these various forms perform and how they evolve. 

An example of Niklas' model showing how simple branching pattern (bottom) can evolve over time into more complex, yet familiar, forms (top).

The most remarkable part of this model is that it inevitably produces all sorts of familiar plant forms, such as those we see in lycophytes, ferns, as well as many of the tree architectural models mentioned above. What's more, later iterations of the model do an amazingly accurate job at predicting forest structure dynamics such as self-thinning, mortality, and realistic size/frequency distributions of various species. 

It would appear that the rules governing what we know as a plant are to some degree universal. Because constraints such as light capture and the passive movement of water are firmly grounded in the laws of physics, it makes sense that the successful plant architectures we know and love today (as well as those present through the long history of plant evolution on this planet) are in large part a result of these physical constraints. It also begs the question of what photosynthetic life would look like on other planets. It is likely that if life arose and made its living in a similar way, familiar "plant" architecture could very well exist on other planets.

Listen to my interview with Dr. Karl Niklas here.

Photo Credits: [1] [2] [3]

Further Reading: [1] [2] [3] [4] [5]

Plants and Music

Turn up the music! My plants can't hear it! Okay, there goes a cheap attempt at humor... In all seriousness, I was always told as a child that plants respond to music. I have since heard many variations on the theme but basically the ideas is that plants, when exposed to music, respond with increased growth. To take things one step further, it would seem that plants have something akin to musical tastes, preferring classical to rock music.

Is there any real scientific evidence to this or is it all just a bunch of silly pseudoscience? Also, if it is true, what could possibly be going on within the plant that causes a response to music, something we thought was reserved to lifeforms with the proper sensory equipment?

The truth is, there is not much real science to base these assumptions on. The internet is full of anecdotal tales and "experiments" that hinge themselves on new age belief systems. In fact, the first "experiments" on how music influences plant growth was done by a woman named Dorothy Retallack. 

Retallack claimed that plants exposed to classical music grew vigorously whereas plants exposed to rock music languished. Considering how much heavy metal my houseplants are exposed to, I think I have more than enough evidence to say otherwise. Besides her poor experimental design, Retallack was heavily motived by quite a conservative, religious agenda. She had it out for mean old rock n' roll and was damned if she couldn't prove her point. What work has been done since Rettalack's time is tantalizing at best but from this point on, keep in mind that the jury is still out on this topic.

So, why would plants respond to music? They don't have ears or anything in their biology that would function as an auditory device, right? Let's re-frame the question in a more basic sense. What is music? Music is nothing more than organized sounds and sounds are nothing more than pressure waves, that is, disturbances in the atmosphere, a process akin to wind. Plants do, in fact, respond to wind, however, wind is a far more physical force than music. Wind can blow over entire swaths of forest whereas music cannot. What mechanism exists that could possibly explain a plant having any kind of response to music? 

Plants respond to heavy wind by growing smaller or by hugging the ground (think alpine vegetation). High winds could generally be seen as a taxing force in the plant world so why would music make plants grow taller and more vigorous? In my opinion, this idea is not a satisfying explanation. As stated above, music doesn't come close to the raw physical power of wind so there could be something else at work. 

In a study done by Margaret E. Collins and John E.K. Foreman out of the University of Western Ontario in London, Canada, they demonstrated that plants responded to different kinds of tones. The tones were either pure (without variation) or random. The results did not show any sort of negative responses from the plants, but rather the plants showed different rates of growth. Plants exposed to pure tones grew better than those exposed to random tones. 

The mechanism they hypothesized for the increased growth in pure tone plants was that the pure tones were able to move air, however slightly, around the leaf. Plants don't like stagnant air and thus, slight air movement is likely to be more beneficial. The random tones did not produce as vigorous of a response, but the plants still grew. It is possible that the random tones caused less air movement around the plants and, because of this, they did not grow as quickly.

Another explanation that seems plausible was put forth by USCB via their science line. They feel that one possible explanation is that the plants aren't the ones responding to the music, but rather the gardener. If you are listening to music while caring for your plants, then chances are it is music you enjoy. If you are like me, then music really has the power to put you in a good mood. If you are in a good mood then chances are you are more likely to take better care of your plants.

All in all, this is an interesting idea. As I said above, the results are mostly controversial and new agey. There are some tantalizing papers that have been published but their methods have been heavily scrutinized. It seems like this is one of the more popular science fair projects for kids to explore and really, anything that gets kids thinking about science and plants is a cool idea in my book. Until more hard science is done on the subject, we can't say for certain. Either way, I will continue to rock out to my favorite tunes and maybe, just maybe, my plants are benefiting from it too.

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