Tag Archives: ecosystem functioning

PSE journal club: Vegetation exerts a greater control on litter decomposition than climate warming in peatlands

A bright, blustery day in England’s north Pennines. Fluffy cotton grass heads bob and bounce on the breeze. On the side of a hill, a strange array of hexagonal, knee-high structures glint and sparkle. Four figures, hunched against the wind, move methodically along jaunty wooden boardwalks, which rest on the blanket bog, crouching at each hexagon in turn. Welcome to Moor House National Nature Reserve, the site of an experiment designed to investigate how a warmer climate will affect the speed with which plant litter is recycled back into the soil, and ultimately the atmosphere.

One of the inherent difficulties associated with upland experiments.

One of the inherent difficulties associated with upland experiments.

If you’re wondering about the shift in tone in the opening paragraph of this #psejclub post compared to some of the others, it’s because I was there. I helped to set up and sample the aforementioned experiment, which is located a few hundred metres away from where I did my PhD fieldwork, in exchange for help with my own work, so I have to admit to having a degree of personal bias! I do think that the work is of general interest, though, and conveys some important findings about litter decomposition in peatlands that will help us to build a picture of how these key processes might change as plant communities shift in response to climate change.

The paper, currently a preprint in ESA Ecology, describes an experiment that uses a combination of open-top, passive warming chambers and plant removal treatments to investigate how the presence of certain plant functional types and warmer temperatures affect rates of litter decomposition. The authors used litter bags, filled with the litter of each plant functional type and buried in plots beneath the plant removal and warming treatments. In all, there were eight treatments (combinations of graminoid, shrub and bryophyte removal, a bare plot and a control with no plants removed), replicated over four blocks, with half the plots warmed by the passive chambers.

So how do peatland processes respond to warming?

So how do peatland processes respond to warming?

The main finding of the paper is that presence or absence of plant functional groups had a stronger effect on peatland litter decomposition than the warming of approximately 1°C achieved by the passive chambers. Removing the shrubs from the peatland resulted in faster decomposition of graminoid and bryophyte litter, after two years. Litter identity was also important – in the first year of the experiment this was the main factor controlling rates of litter decomposition, with bryophyte litter decomposing most slowly, followed by shrubs and graminoids. After two years, the live plants present in the plot (i.e. presence of shrubs) were more important than the litter identity. Warming affected the composition of the bacterial community, while the fungi responded more strongly to the presence of shrubs.

While these results are compelling, they should be taken with caution, as the authors suggest: since the duration of the experiment was two years, further interesting effects resulting from the decomposition of shrub and bryophyte litter, which happens more slowly than in graminoids, might not have been captured. After four or five years, the decomposition of more recalcitrant litter could reveal more interesting effects. The same is true for the warming treatment: given a longer study period, the effect of 1°C warming on plant litter decomposition might become more important. It is, however, easy to write ‘more long-term experiments needed!’ while, in reality, the amount of effort required to maintain the plant removal treatments and warming chambers in the harsh upland environment of the Pennines represents a considerable hurdle. And one has to motivate one’s volunteers to see past the inherent absurdity of weeding a moorland!

Overall then, what’s the message? Dead or alive, whether you’re graminoid, shrub or bryophyte seems to exert much stronger control on litter decomposition rates in peatlands than temperature. While warming doesn’t have much of an effect on plant decomposition, it does affect the bacterial community in the peat, which might have important implications for graminoid decomposition, since bacteria are well-equipped to munch through labile substrates. Fungi respond when you take away the shrubs, which provide the more recalcitrant litter they’re specialised for dealing with, and might therefore moderate the response of the peatland carbon cycle to warming. Given a longer time period, more effects may emerge from this experiment.

I’m interested to know about the responses of decomposition processes in other ecosystems to warming and plant functional group removal. In grasslands, for instance, what happens when you increase the ambient temperature and remove the nitrogen fixers? Of course, these sorts of studies require a sufficiently long period to become stable and start producing results. In a time of apparently perpetually shrinking budgets, what’s next for long-term field experiments in ecology and biogeochemistry? What are the current barriers to their deployment, or do we in fact have enough to answer our more pertinent questions? Let me know what you think – get involved by commenting on this post, and posting on Twitter and Facebook using the #psejclub tag. I’m looking forward to hearing from you!

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Soil biodiversity and soil community composition determine ecosystem multifunctionality

This paper, by Cameron Wagg et al., which was published online early in PNAS last month, describes the results of a very interesting experiment in which the authors manipulated soil biodiversity and measured the effect of these manipulations on a range of ecosystem functions.

More specifically, they created a gradient of reduced soil biodiversity (including a range of faunal and microbial groups) by sieving the soil through a number of decreasing mesh sizes, adding the fraction that passed through the sieve to sterilized soil, while also adding the sterilized fraction that remained on top of the sieve. They then grew plant communities consisting of common grasslands species in the soil for 14 and for 24 weeks, in two separate experiments. At the end of the first experiment, and after 12 and 24 weeks of the second experiment, they measured plant diversity and productivity, carbon sequestration, litter decomposition, nitrogen turnover, N2O emission, phosphorus and nitrogen leaching as ecosystem functions, and fungal and bacterial diversity (by TRFLP), mycorrhizal root colonization (microscopically), and nematode abundance (microscopically).

They then used these data to relate the ecosystem functions measured to the soil biodiversity treatments. In addition, they calculated z-scores for the range of ecosystem functions measured as well as for all groups of organisms quantified, and regressed these against each other to answer the question whether ecosystem multifunctionality is related to soil biodiversity. This approach, of summarising a number of ecosystem processes into one ecosystem multifunctionality index, has been used previously by Maestre et al. (2012).

Their findings are very interesting and will make a lot of soil ecologists very happy: they find that a number of the individual ecosystem functions are reduced with declining biodiversity, but also that ecosystem multifunctionality is positively correlated with overall soil biodiversity.

When taking a closer look at the data, it becomes clear that the reduction in soil biodiversity varies between groups and isn’t linear with the decreasing mesh sizes – mycorrhiza and nematodes drop down sharply after the third ‘dilution’, whereas the other parameters show a more gradual decline. The authors have taken this into account by not only relating ecosystem functioning to the diversity treatments, but also to the abundance and diversity of individual groups. When taking a closer look at this, it becomes clear that the microbial properties measured have a far stronger effect than nematode abundance. In addition, the effect of reduced soil biodiversity on a range of functions is indirect, through effects of plant productivity and diversity.

Of course, it is very easy to criticise aspects of this study. You can question whether bacterial and fungal diversity, microbial biomass, mycorrhizal colonization, and nematode abundance together are a realistic representation of soil biodiversity. For example, why was nematode diversity not assessed? And why not higher trophic levels, such as Collembola and mites? Microbes and nematodes are only a fraction of the soil food web (Fig. 1). With the current analyses, the title ‘Soil microbial diversity and community composition determine ecosystem multifunctionality’ might have been more appropriate.

A (simplified) example of a soil food web, with the groups measured by Wagg et al. (2014) indicated by the dashed line.

A (simplified) example of a soil food web, with the groups measured by Wagg et al. (2014) indicated by the dashed line.

Also, it would have been interesting to see root biomass in addition to mycorrhizal colonisation – a number of recent papers point to the importance of roots for ecosystem functioning (e.g. Orwin et al. 2010, Grigulis et al. 2013)

A more technical comment relates to the measurement of nitrogen turnover – this was assessed by measuring the uptake of 15N from Lolium multiflorum litter into aboveground L. multiflorum biomass. So, this measurement might be a proxy for L. multiflorum biomass, which decreases with decreasing soil biodiversity, rather than for nitrogen turnover.

On another note, and I would be very interested in other people’s opinion, I am wondering about the value of using an index for ecosystem multifunctionality. True, this averages across ecosystem functions and can therefore inform management to optimize overall ecosystem functioning. However, are the ecosystems that have the greatest average functioning really the most sustainable, and thus, desirable ecosystems? Are all ecosystem functions equally important? There might be trade-offs between different ecosystem functions – for example between crop yield and nitrogen retention, or between decomposition and carbon sequestration. We might want to optimize a certain function in a certain area, of which we already know that it has potential in delivering a certain function, rather than promoting multifunctionality across the board. For example, peatlands store large amounts of carbon because of their low decomposition rates, and agricultural production systems have high yields but low carbon sequestration.

However, in this paper, the multifunctionality index serves the purpose of summarizing overall ecosystem functioning, which shows a strong and positive relationship with soil biodiversity. Done like this, it summarizes a range of measurements that non-specialists might struggle to interpret – thus, it simplifies and reinforces the message of the paper that soil biodiversity determines ecosystem functioning.

Experiments like this require an enormous amount of work, and you simply can’t include everything. It is incredibly difficult to modify soil biodiversity without simultaneously changing soil properties, and the authors of this paper have achieved this by used an elegant method of reducing soil biodiversity. Thus, in contrast to many earlier studies, they were truly able to mechanistically elucidate the role of groups of soil organisms in ecosystem functioning.

This paper adds to the growing body of literature that soil biodiversity plays a crucial role in ecosystem functioning, and highlights the importance of conserving, and promoting, soil biodiversity. That’s what I like to hear!