For real guys, bigger isn't always better

In my previous post I alluded to a study that framed a potential answer to the question: Why are there so many small plants? Well, here is that study-its results are fascinating.

As I mentioned before, traditionally people assume that bigger is better in terms of plant competition. I also mentioned that the majority of plants are small. So, if the majority of plants are small, how come most plants aren’t big if that is considered to be so advantageous? 
 

It is known that bigger plants generally produce more viable offspring- and this makes sense, right? They usually have more flowers and thus more fruits and more seeds. They are also less sensitive to some of the physical constraints of the environment. So, in the “traditional plant competition theory boxing ring” round 1 goes to the big guys. Let’s think about this though. We have a few big guys with a lot of offspring. But we have way more small guys than big guys. So does the collective offspring of the small guys trump the big ones? Round 2 here we come.

Jesse Chambers conducted a natural field experiment to test this idea. She sampled natural populations of 21 herbaceous angiosperm plant species throughout the Kingston area. Each population was harvested at reproductive maturity by placing a 1x1m plot in the area of highest density-it was assumed this would be where crowding was the most intense.  Within the 1x1m plot, reproductive plants were harvested, bagged, dried and weighed. Plant size frequency distributions were created by dividing the range of individual plant sizes for each population sample into 10 equal deciles of plant size. Relative reproductive output per size decile was calculated as the relative total mass per decile relative to the grand mass total for the entire population. Estimated reproductive output was generated under the assumption that total seed production is proportional to plant mass. 

Figure 1: Size and fecundity relationships for Cardamine parviflora

Here is what Jesse found- it’s pretty cool, but expected. All species had size distributions that were strongly right-skewed. The number of seeds per plant was counted for the entire population of the species Cardamine parviflora and it was found that the mean seed mass did not differ between the 10 largest and smallest plants and fecundity was directly proportional to plant size (See Figure 1). This confirmed the estimate of relative reproductive output. 

Here’s the kicker: For each of the 21 herbaceous study species, the vast majority of the offspring production within the population was contributed by the three, four, or five smallest deciles of plant size. And in 7 of those species the majority was from the 2 smallest deciles of plant size distribution. 

Wait so the smallest plants are collectively producing the most offspring in crowded vegetation? Yeah! They are!

So what does this all mean then? Well, as it says in Chambers and Aarssen “A renowned chemical evolutionist, Leslie Orgel is credited with saying, “Evolution is cleverer than you are””. It seems, well, it seemed intuitive that larger plants would have the ability to produce large and highly fecund offspring, a huge component of plant fitness and that this made bigger better. Chamber’s study is interesting in that is opens the eyes of plant ecologists to a new potentially important component of plant fitness- that being the ability to produce offspring that will survive AND reproduce before death despite having to live as small, suppressed weaklings. This is evidence for the concept of plants having reproductive economy. Reproductive economy is a product of natural selection in plants as a result of the inability of most plants to escape being small as a result of crowding. 

So, bigger isn’t necessarily better. Natural selection does favour large plant size, just not most of the time. Most of the time, plant communities and populations are crowded where most plants are relatively the same, suppressed size. 

I think it’s a knock-out. Round 2 goes to the small plants!





Source: Chamber, J and Aarssen, L.W. 2008. Offspring for the next generation: most are produced by small plants within herbaceous populations. Evolutionary Ecology. 23:737-751.

Consider this.

Consider this:

 According to traditional plant competition theory, bigger is better. Plants with a large body size are better at capturing resources and space from their neighbours and they have a clear advantage in the race towards the canopy. The bigger plant would be a better competitor and would likely have a higher fitness.

The above ideas have been part of the underlying focus of much of the research done in the Aarssen lab to date. But what makes that so interesting? Why do we care? 

Consider this:

Plant size distribution is right-skewed at virtually all scales. This means that the majority of plants have a relatively small plant body size. 

Wait...so it’s generally accepted in the literature that bigger is better. But if bigger really was better...why then are there so many small plants? 

Consider this yourself. Post some comments with potential explanations for this paradox. 

My next post will discuss a beautiful, natural experiment done in the Aarssen lab, which provides a mind-blowing potential explanation for this paradox.

Precision, Generality and Realism

It is nearing the end of the second week of classes and I am in the process of designing a project for my M.Sc. thesis. A couple of days ago I met with my supervisor to discuss some potential ideas for my project. It is not uncommon for him to rant about all sorts of different ideas; however, this time something he said really sparked a fire in me. He pointed out that my B.Sc.H thesis was a natural experiment that involved no set-up or manipulations of any kind and that he really appreciates the beauty of these “realist” experiments- an experiment where you simply go out and record or collect exactly what you see. This inspired me to do a literature review of this idea of natural experiments and I came across a very interesting paper in which Richard Levins addresses different types of biological studies.

In 1966, Levins wrote a paper titled Strategy of Model Building in Population Biology published in American Scientist. He explained that it is ideal to work with mathematical models in biology that maximize three things: 1) Generality, 2) Realism and 3) Precision, but that it is never possible to maximize these three things simultaneously.

Levins suggests that there is a trade-off between these three ideals and that three strategies have evolved to deal with this problem. For example, one could sacrifice generality to realism and precision. The focus of an experiment could be on the behaviour of an organism within very small parameters that would yield real and precise results but would tell us very little about the organism’s behaviour in general. Scientists could also sacrifice realism to generality and precision in which general equations and models can be developed that yield precise results but are extremely unrealistic. Finally one can sacrifice precision to realism and generality which rely on flexible models that assume wide variance in different functions.

Different scientists prefer different methods and each way has its merit. In my blog, I will discuss current studies in ecology that sacrifice generality or precision for realism and why I believe this approach is the most valuable.

Source: Levins, Richard. 1966. The Strategy of Model Building in Population Biology. American Scientist 54(4): 421-431.