Wednesday, April 25, 2018

Don't forget the details! Trait ecology and generality

The search for generality is perhaps the greatest driver of modern ecology and probably also the greatest source of ecological angst. Though ecological trends frequently reflect the newest, brightest hope for generality, the search for generality (perhaps by definition) encourages us to ignored details and complexities. Maybe this means that some areas of study won't develop fully until they've fallen out of fashion. And maybe this means that the most interesting science happens when the pressure to 'save community ecology' is gone. A great example of the kind of post-hype, thoughtful approach for trait-based ecology comes from Reynolds et al. (2017) in Tree Physiology. They do a really nice job of highlighting some of the details that must inform trait-based ecology. Here, Reynolds et al. take a broad comparative approach across species, but incorporate important details that have at times been overlooked - especially the role of the environment, recognizing and measuring both constitutive and plastic traits, captures that there are multiple paths (or trait combinations) that can result in similar functioning.

The authors look at four conspecific tree species (Brachychiton spp.) with different average positions along an observed moisture gradient (CMD or climate moisture deficit). Two species occupied drier areas of Australia ('xeric species'), while the other two were found in more moderate areas ('mesic species'). The authors assumed that the different distributions of these species reflect different hydraulic niches. Were species' hydraulic niches associate meaningfully with their traits, specifically those trait associated with drought stress responses. Though these species are closely related--and so huge divergences in form and function might not be expected--the costs and benefits of drought resistance should differ among the species. In dry environments, drought resistance strategies should be more important, and may select for particular traits or sets of traits. Trait states associated with drought conditions include "reduced leaf area, enhanced stomatal control, safer or more efficient xylem, increased tissue water capacitance...and/or deeper root systems " may all be selected for. On the other hand, investment in these traits when water is not limited is often costly, reducing growth and competition. This suggests a meaningful selective regime associated with the CMD gradient and trait values might exist.

One important, but oft-overlooked aspect of trait ecology is that trait values depend on both genes and the environment. Reynold et al. incorporate this fact this by manipulating water availability between drought and control treatments. They measured both constitutive components of trait values – those driven by genetics and expressed regardless of environments – and the plastic or environment-dependent components. For instance, in the presence of prolonged drought, trees might increase root production or change leaf characteristics. In addition to manipulating water availability between treatments, the authors measured nine traits related to morphology and allocation.
From Reynolds et al. 

Given the expectation that trait values reflect the complex interaction of genetics and the environment in different species, is it possible to even make simple predictions about trait-environment relationships? The authors find that "These complex relationships illustrate that assuming that individual traits (often measured on individuals under a single set of environmental conditions) reflect drought resistance is likely to be overly simplistic and may be erroneous for many species. However, our results do suggest that generalization may be possible, provided multiple traits are measured to explore specific integrated drought strategies."

Indeed, some results are relatively predictable relationships: under well-watered control conditions, the allocation of biomass matched the expectation: xeric species had higher investment in below-ground biomass and in transport tissues than the mesic species (both characteristic of a water-conserving species).

On the other hand, leaf traits such as SLA did not show any trend related to species' assumed drought tolerance, either for constitutive or plastic trait components. Sometimes traits associated with the leaf economic spectrum such as SLA are assumed to indicate stress tolerance, but this was not the case.


By far the most interesting result was the observation that the xeric species had the highest assimilation and stomatal conductance rate and the lowest water use efficiency under well-watered conditions. Only by also examining these species under drought conditions was it possible to observe that they are highly plastic with regards to water use efficiency. In fact, they show a feast or famine approach to water usage - "where high photosynthetic rates per unit leaf area and high investment in root and stem tissue even in well-watered conditions are achieved through profligate water use during rare periods of water availability, in order to establish a root system and stem storage tissues necessary to survive long periods of water stress." Under drought conditions, these species show reduced root tissue investment; in contrast, mesic species follow expected patterns and plastically increase root tissue investment.

This paper is a reminder that the details are also fascinating and informative. As humans, we may have a simplistic understanding of the realized environment sometimes. To us perhaps all water stress is similar, but for each species in this study, the long term selective pressures may be meaningfully different - in timing, duration, and life stage. This creates the potential for complex differences between species which may best be reflected via life history strategies involving multiple traits. That may still imply some degree of generality is possible, but it is multi-dimensional.

Works cited:
Victoria A Reynolds, Leander D L Anderegg, Xingwen Loy, Janneke HilleRisLambers, Margaret M Mayfield; Unexpected drought resistance strategies in seedlings of four Brachychiton species, Tree Physiology, https://doi.org/10.1093/treephys/tpx143

Wednesday, April 4, 2018

Life in Plastic Ain’t so Fantastic

Guest post by Louis Vassos, MEnvSci Candidate in the Professional Masters of Environmental Science program at the University of Toronto-Scarborough


Much like the Buggles’ 1980 debut album, our material preferences are well within the age of plastic. Thanks to its light weight, durability, inertness, and low manufacturing costs, our use of plastics has increased dramatically since the mid-20th century. From bottles and toys to car parts and electronics, there is seemingly no application beyond its reach. Despite its uses and benefits, it has come under increasing scrutiny by environmentalists in recent years. In this regard, we tend to think of larger-scale and more visible environmental impacts, such as accumulation in landfills and petrochemical use in manufacturing. There has also been a significant amount of research on plastic in marine environments, usually focused on larger debris known as macroplastics. Over the past decade, however, there has been increasing concern about a new type of plastic debris in our oceans. Though its presence was first highlighted in the 1970s, we are only just beginning to realize the impact of fragments known as microplastics. As their name would suggest, they are small pieces of plastic, typically measuring less than 5mm in diameter and sorted into two distinct classifications.


Primary microplastics are manufactured to be microscopically sized and are typically used in air blasting as a paint and rust remover, as well as in personal care products as an exfoliating scrubber. This latter use has risen sharply in cosmetics and facial cleansers since the 1980s, with plastic “microbeads” replacing natural materials such as pumice and ground almonds. Regardless of application they usually enter water bodies through drainage systems, and are easily able to pass through filtration systems at sewage treatment plants due to their small size.

Microbeads in toothpaste. Retrieved from: https://blog.nationalgeographic.org/2016/04/04/pesky-plastic-the-true-harm-of-microplastics-in-the-oceans/




Secondary microplastics arise from the breakdown of larger pieces of plastic debris on both land and in water. Larger debris will typically enter marine ecosystems directly or indirectly through careless waste disposal, often being transported through river systems. Sources of transfer include coastal tourism, extreme weather events, fishing, other marine industries, and accidental spillage during transportation. Over time, a culmination of processes such as exposure to UV radiation can reduce the debris’ structural integrity, causing brittleness, cracking, and yellowing. This in turn can lead to fragmentation through abrasion and waves, and fragments will gradually become smaller over time before reaching microplastic size (Cole et al, 2011).


As Eriksen et al (2014) have estimated, there is a minimum of 5.25 trillion plastic particles weighing 268,940 tons in the world’s oceans. Microplastics account for 92.4% of this mass, and their reach has been substantial. Because of their buoyancy and durability, they have the ability to travel long distances without degrading for years. Denser plastics (such as PVC) will sink and have the potential to reach coastal sediment (Andray, 2011). Other marine microplastics will end up trapped in ocean current systems known as gyres, the most famous grouping of which is the “Great Pacific Garbage Patch” in the North Pacific Gyre. Despite what the name would suggest, it is not an island-like mass of floating debris, but is more akin to an extensive “soup” of debris difficult to see with the naked eye. At a density of 334,271 pieces/km2, microplastic mass in the area was found to be 6 times that of plankton (Moore et al, 2001). 
Potential microplastic transport pathways (From Wright et al, 2013)

Densities such as this increase potential microplastic ingestion by various marine organisms, especially filter feeders, plankton, and suspension feeders. These species may mistake debris for prey based on size or colour, or passively ingest them without being selective (Wright et al, 2013). In Farrell and Nelson’s (2013) study of mussel-eating crabs, they found that it is possible for microplastics to be transferred to individuals at a higher trophic level. The large surface area to volume ratio of microplastics makes them susceptible to water-borne pollutant contamination, and can cause toxic plastic additives such as BPA and PCB to leach into the water. This debris can also act as a dispersal vector for microbial communities, including potentially pathogenic species (Jiang et al, 2018). While the ingested debris can accumulate within individuals and be transferred up the food chain, the exact effects of this are not entirely known at this point in time (Avio et al, 2017). A recent study by Lei et al (2018), however, found that microplastics can cause oxidative stress and intestinal damage in zebrafish and nematodes, and that their toxicity is closely dependent on particle size.
Intestinal damage in zebrafish caused by exposure to 1.0 mg L-1 of different microplastic types and sizes. Photograph A shows control (top), survival (middle), and dead after exposure (bottom) zebrafish (From Jiang et al, 2018)
Fluorescent microspheres on a crab’s gill lamella transferred from ingesting mussels, each measuring 5 micrometres in diameter (From Farrell and Nelson, 2013)

          What does the future hold for microplastics? Because their effects on both marine life and humans is relatively unknown, it is important to try and prevent them from entering and accumulating within marine environments. Properly dispose of larger plastic items to prevent them from entering waterways and breaking down into secondary microplastics, and be conscious about the presence of primary microplastics in other products. Make informed decisions when buying cosmetics, and choose ones that use natural exfoliating materials. Microbead bans have already begun to be enacted in several countries, including the UK, US, Canada and New Zealand (Pfeifer, 2018). There is also the potential for future studies on topics such as the health effects of microplastic ingestion and leached additives, debris behavior within the water column, and new standardized techniques for detection and sampling (Cole et al, 2011). It is hard to say what will happen next, but the removal of these 5.25 trillion particles from our oceans will prove to be a very difficult challenge without the development of novel extraction methods.



SOURCES

Anadrady, A.L. 2011. Microplastics in the marine environment. Marine Pollution Bulletin 62:1596 – 1605
Avio, C.G., S. Gorbi, and F. Regoli. 2017. Plastics and microplastics in oceans: from emerging pollutants to emerged threat. Environmental Research 128: 2 – 11
Cole, M., P. Lindeque, C. Halsband, and T.S. Galloway. 2011. Microplastics as contaminants in the marine environment: a review. Marine Pollution Bulletin 62:2588 – 2597
Eriksen, M., L.C.M. Lebreton, H.S. Carson, M. Thiel, C.J. Moore, J.C. Borerro. F. Galgani, P.G. Ryan, and J. Reisser. 2014. Plastic pollution in the world’s oceans: more than 5 trillion plastic pieces weighing over 250,000 tons afloat at sea. PLOS One
Farrell, P., and K. Nelson. 2013. Trophic level transfer of microplastic: Mytilus edulis (L.) to Carcinus maenas (L.). Environmental Pollution 177:1 – 3
Jiang, P., S. Zhao, L. Zhu, and L. Daoji. 2018. Microplastic-associated bacterial assemblages in the intertidal zone of the Yangtze Estuary. Science of the Total Environment 624:48 – 54
Lei, L., S. Wu, S. Lu, M. Liu, Y. Song, Z. Fu, H Shi, K. Raley-Susman, and D. He. 2018. Microplastic particles cause intestinal damage and other adverse effects in zebrafish Danio rerio and nematode Caenorhabditis elegans. Science of the Total Environment 619:1 – 8
Moore, C.J., S.L. Moore, M.K. Leecaster, and S.B. Weisberg. 2001. A comparison of plastic and plankton in the North Pacific Central Gyre. Marine Pollution Bulletin 42:1297 – 1300
Pfeifer, H. 2018. The UK now has one of the world’s toughest microbead bans. CNN. Retrieved from: https://www.cnn.com/2018/01/09/health/microbead-ban-uk-intl/index.html
Wright, S.L., R.C. Thompson, and T.S. Galloway. 2013. The physical impacts of microplastics on marine organisms: a review. Environmental Pollution 178:483 – 492