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Effects of Microplastic Fibers and Drought on Plant Communities

Yudi M. Lozano* and Matthias C. Rillig
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ABSTRACT: Microplastics in soils can affect plant performance, as shown in studies using
individual plants. However, we currently have no information about potential effects on plant
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community productivity and structure. In a plant community consisting of seven plant species
that co-occur in temperate grassland ecosystems, we thus investigated the effect of microplastics
(i.e., microfibers) and drought, a factor with which microfibers might interact, on plant
productivity and community structure. Our results showed that at the community level, shoot
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and root mass decreased with drought but increased with microfibers, an effect likely linked to
reduced soil bulk density, improved aeration, and better penetration of roots in the soil.
Additionally, we observed that microfibers affected plant community structure. Species such as
Calamagrostis, invasive in Europe, and the allelophatic Hieracium, became more dominant with
microfibers, while species that potentially have the ability to facilitate the establishment of other
plant species (e.g., Holcus), decreased in biomass. As microfibers affect plant species
dominance, the examination of cascade effects on ecosystem functions should be a high priority
for future research.

■ INTRODUCTION
Microplastics, a diverse group of polymer-based particles (<5
films or Lolium perenne (grass) exposed to fibers had shown
reduced biomass6,13 while Allium f istulosum (crop) exposed to
mm), are becoming recognized as an important new global fibers had shown an opposite effect.14 Likewise, Plantago
change factor potentially influencing terrestrial ecosystems.1,2 lanceolata (forb) and Allium f istulosum14,15 had contrasting
Between 1950 and 2015, global plastic waste is estimated to responses to microplastics in relation to root morphological
traits (e.g., root length). This variety of individual plant
have been 6300 million tonnes.3 The abrasion of large plastic
responses suggests that different plant species in a community
objects during manufacture or in the environment, the erosion
could be affected to a different degree by the addition of
of tires when driving, or the abrasion of synthetic textiles
microplastics in the soil, which could potentially affect plant
during washing4 are just some of the sources of large amounts
productivity and community structure.16 Therefore, in a
of microplastics that will end up not only in the oceans but also
community, some species may be better able to take advantage
in terrestrial systems.
of the changes in soil properties due to the presence of
Microplastics can enter soils via soil amendments,1,5 plastic
microplastics.
mulching, irrigation,6,7 diffuse urban runoff, flooding, and
Depending on the microplastic effects on different plant
atmospheric fallout.8,9 As consequence, microplastics in soil
species in any given community (e.g., invasive or facilitative
may appear in the form of fibers, films, granules, presenting a
species), different ecosystems functions could be affected. Dry
variety of shape, composition, and abundance in concen-
grasslands species show different responses to drought through
trations that may reach up to 7% close to industrial areas.10
the adjustment of their root morphological traits;17,18 there-
Therefore, microplastic effects on terrestrial systems depend
fore, microfibers could affect root traits as well, as a
on the microplastic type and the plant−soil system involved.
consequence of their effects on soil-water dynamics,14 which
For instance, agriculture lands would be highly affected by
at the end, would modify resource availability affecting plant
microplastic films6,11 as plastic mulching is intensively used to
competition.19 Previous research provides evidence that
promote plant productivity,11 while rivers or roadsides would
grasses and herbs may have contrasting strategies to face
tend to be more affected by microplastic fragments or
drought.17 For example, Festuca and Holcus respond with
microfibers.9,12 Likewise, microplastic effects in terrestrial
systems may be influenced by the soil type (e.g., sandy vs
loamy soil) and by the plant species, as depending on their Received: February 19, 2020
identity, they respond differently to environmental stressors. Revised: April 14, 2020
The effects of microplastics on plant performance have been Accepted: April 14, 2020
observed in some species, such as crops or plants growing as Published: April 14, 2020
single individuals or in a population but not in a community
context. For example, Triticum aestivum (wheat) exposed to

© 2020 American Chemical Society https://dx.doi.org/10.1021/acs.est.0c01051

6166 Environ. Sci. Technol. 2020, 54, 6166−6173
Environmental Science & Technology
increased specific root length (i.e., root finesses), likely
promoting mycotrophy for resource acquisition, while herbs
such as Achillea or Potentilla respond with decreased specific
root length but instead increase root biomass, likely as a
strategy to promote water acquisition.17,20 Thus, we could
expect that at the community level, root morphological traits
may be affected by drought, an effect likely intensified by
microfibers in the soil. Additionally, we could expect a better
performance of fast growing species (e.g., most grasses), as
they could rapidly access the limited water.21 Several models
suggest that grasses and herbs could have complementary
resource use under stressful conditions,22,23 which suggest that
microplastics exacerbating the effects of drought may influence
species abundance.
Among microplastics, microfibers are considered one of the
most predominant microplastic types in the soil.7,8 The linear
shape, size, and flexibility of such particles could reduce soil
bulk density and soil aggregation,14 although they can also
entangle soil particles and thus contribute to the soil aggregate
formation process16 so that one of the most critical effects of
microplastic fibers is on soil structure and soil-water dynamics.
Microplastic fibers can enhance water holding capacity and
keep water saturation for longer periods,14 but it is possible
that they increase soil water evaporation as additional channels
for water movement may be created.24 Changes in soil-water
dynamics could alleviate or exacerbate drought, a phenomenon
which is predicted to increase over the next few decades25 in
many regions worldwide, with the consequences on plant
communities structure and ecosystem functionality remaining
unknown.
In this study, we established a community of grasses and
herbs including plants with invasive, allelopathic, or facilitative
characteristics, which naturally co-occur in temperate dry
grasslands in northeastern Germany. We assessed the effect of
microfibers in the soil and drought on plant productivity and
structure. We hypothesized that microfibers and drought
would strongly alter not only productivity but also community
composition and that microfibers may exacerbate the effects of
drought on plant communities.
■ MATERIALS AND METHODS
Species Selection. We selected seven plant species,
including grasses such as Festuca brevipila, Holcus lanatus,
and Calamagrostis epigejos and herbs such as Achillea
millefolium, Hieracium pilosella, Plantago lanceolata, and
Potentilla argentea to shape community modules, typical of
temperate grasslands ecosystems. Seeds of these plant species
were obtained from a commercial supplier in the region
(Rieger-Hofmann GmbH, Blaufelden, Germany). We will refer
to plant species by their generic names from now on. All are
common, frequently co-occurring grassland species in Central
Europe that naturally grow in the same patch as seen in field
observations in dry grasslands in the Brandenburg region
(Germany). As microplastic fibers could alleviate or exacerbate
drought, we selected plant species that exhibit contrasting
responses to drought conditions. Growing as single species,
Festuca and Holcus have decreased root biomass with drought,
while Achillea, Plantago, or Hieracium show the opposite trend.
Other traits, such as root diameter or root finesses (SRL), also
differ among the selected species as drought increases.17
Microplastic Fibers. Polyester fibers (Rope Paraloc
Mamutec polyester white, item number, 8442172, Horn-
bach.de) were manually cut with scissors, and a length of 5.0
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mm was established as an upper size threshold in order to
generate microplastic fibers. They had a length of 1.28 ± 0.03
mm and a diameter of ∼30 μm. Size distribution is given in
Figure S1, and mechanical properties of polyester fibers are
listed in Table S1.
Soil Preparation. We collected sandy loam soil (0.07% N,
0.77% C, pH 6.66) from Dedelow, Brandenburg, Germany
(53° 37′ N, 13° 77′ W) where our plant species naturally grow.
The soil was dry when collected, and then it was sieved (4 mm
mesh size), homogenized, and mixed with microplastic fibers at
a concentration of 0.4%. A 12 g portion of microfibers
(∼763333 fibers g−1 microplastic) was mixed into 3 kg of soil
for each pot. Soil preparation was done separately for each
experimental unit: microplastic fibers were separated manually
and mixed with the soil for 30 min in a large container, before
being placed into each individual pot, to help provide an equal
distribution of microfibers throughout the soil (optical
inspection) and the correct microfiber concentration. This
was done in order to reflect the fact that after accumulation at
the top of the soil, microplastics can move along soil profile
due to soil biota activity, properties of the soil particles, and
plant processes such as root growth.26,27 Twenty experimental
units (pots) were established. Half had soil with microfibers,
while the other half had soil without added microfibers. Soil
was mixed in all experimental units in order to provide exactly
the same disturbance.
Experimental design. In May 2019, we established the
experiment in a controlled glasshouse with a daylight period
set at 12 h, 50 klx, and a temperature regime at 22/18 °C day/
night with a relative humidity of ∼40%. Prior to germination,
∼200 seeds per species were surface-sterilized with 10%
sodium hypochlorite for 5 min and 75% ethanol for 2 min and
then thoroughly rinsed with sterile water. Then, the seeds were
germinated in trays with sterile sand and individual seedlings of
similar size were transplanted into pots (16 cm diameter, 16.5
cm height, 3L) 3 days after germination. Twenty-one holes
were dug in a grid, keeping a distance of 2.5 cm among them
(Figure S2). Seedlings were randomly distributed, and each
planting hole received one plant individual, so that a
community module shaped by three individuals of each of
the seven plant species was established per pot. Pots were well-
watered (100 mL twice a week) during the first 3 weeks of
growth. Then, half of them were kept at approximately 70% of
soil water holding capacity (WHC) by adding 200 mL of
water, while the other half were kept at approximately 30%
WHC by adding 50 mL of water. By using 30% WHC, the
plant species we selected, were under stress conditions due to
drought.17,18 We determined the respective amount of water
for our soil type. Pots were watered by hand twice a week for
two months by gently spraying distilled water on the soil
surface. We thus had 20 experimental units in a fully crossed
orthogonal design that includes two microfiber treatments
(one with and the other without added microfibers, also called
“present” and “absent”) and two drought treatments (with and
without drought, also called “drought” and “well-watered”),
with five replicates each (n = 5). All pots were randomly
distributed in the chamber, and their positions were shifted
twice to homogenize environmental conditions during the
experiment. Pots were weighed weekly to verify maintenance
of their respective water content. No changes in the watering
amount or frequency was needed. Likewise, no water drained
out from the pots, so that microfibers added to the soil did not
leave the pots. The percolation behavior of the fibers is
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Table 1. Results from Linear Models on Plant Community and Root Morphological Traits Response to Microplastic Fibers
(i.e., Microfibers, M), Drought (D), and Their Interaction (M x D)a
Plant community traits Root morphological traits
df Shoot mass Root mass Root:shoot Evenness RAD RTD SRL SRSA
Microfibers (M) 1 4.25 (0.05) 11.20 (0.004) 7.67 (<0.01) 5.44 (0.03) 1.88 (0.18) 0.43 (0.52) 1.80(0.19) 1.83 (0.19)
Drought (D) 1 1524.6 (<0.01) 19.52 (<0.01) 1.43 (0.24) 0.32 (0.57) 12.7(<0.01) 14.04(<0.01) 2.49(0.13) 0.006(0.9)
MxD 1 0.22 (0.64) 0.20 (0.65) 2.00 (0.17) 1.19 (0.29) 0.001 (0.96) 1.48 (0.24) 0.03(0.86) 0.36 (0.55)
a
Community evenness is based on species biomass. Root morphological traits: root diameter (RAD), root tissue density (RTD), specific root
length (SRL), and specific root surface area (SRSA). F and p-values (in parentheses) are shown. All significant values (<0.05) in bold.
Heteroscedasticity was corrected for shoot and root mass in the microfiber treatment, evenness in the water treatment, and root:shoot in both.

Figure 1. Microplastic fibers and drought effects on (A) shoot mass, (B) root mass, (C) root:shoot, and (D) community evenness. The box in each
boxplot shows the lower, median, and upper quartile values, and the whiskers show the range of the variation. Data points are shown as circles; n =
5.

unknown, but given their linear shape, it would be expected to
be much slower compared to that of beads. At harvest, shoot
mass was sorted by plant species. Roots were carefully removed
from the soil and gently washed (roots could not be separated
by species). Morphological traits were determined for fine
roots (i.e., <2 mm in diameter): length, surface area, volume,
and root average diameter were measured on a fresh sample of
the plant community by scanning the roots using the
WinRhizoTM scanner-based system (v.2007; Regent Instru-
ments Inc., Quebec, Canada). Then, roots were dried at 60 °C
for 72 h as some trait calculations are based on their dry
weight: root tissue density (RTD; mg cm−3), specific root
length (SRL; cm mg−1), and specific root surface area (SRSA;
cm2 mg−1). Root and shoot masses were measured after the
samples were dried at 60 °C for 72 h. In additional pots with
and without added microfibers, we extracted a soil core and
measured its volume to obtain soil bulk density (n = 4).
Statistical Analyses. The experiment had a fully crossed
orthogonal design where microfibers, drought, and the
interaction were considered as fixed factors. Community root
traits: root average diameter (RAD), root tissue density
(RTD), specific root length (SRL), and specific root surface
area (SRSA), along with community shoot and root mass,
shoot mass per species, and soil bulk density were analyzed
using general linear models. Shoot mass per species accounted
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for the shoot mass of the neighbors (the other six plant
species) as a covariate. Community root:shoot ratio included
shoot mass as the sum of all plant species per pot. Soil bulk
density was examined using microfibers as factor. A general
linear model is a regression model that assumes normality,
homogeneity, and independence of the residuals. Model
residuals were checked to validate these assumptions.28
When necessary, we implemented the varIdent() function to
account for heterogeneity in the microfiber and drought
treatment, respectively (see details in the tables). Principal
component analysis (PCA) of the shoot mass per species was
performed using the function “prcomp” and “fviz_pca” from
the package “factoextra”.29 Ellipses in the PCA graph grouped
the different treatments with a confidence level of 0.95. To
analyze changes in plant community composition, in terms of
biomass, we log-transformed biomass per species and used the
function Manova from the package “MASS”.30 We tested the
null hypothesis by means of the Pillai trace statistic, which
accounts for the variance between groups and is robust to
violation of multivariate normality and homogeneity of the
variance-covariance matrix. Manova is based on the Mahala-
nobis distance which makes it not only appropriate but even
advantageous in terms of statistical power in our case where
there are no zeros in species biomass, i.e., no richness
effects.31,32 Community evenness or equitability describes how
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Figure 2. Principal component analysis of the plant community structure (based on species biomass) growing under well-watered and drought
conditions with or without microplastic fibers (microfibers) in the soil; n = 20. Arrows indicate plant species, which are designated by their genus
names.

Figure 3. Shoot mass (g) of plant species growing in a community under drought and well-watered conditions with or without microplastic fibers
in the soil. Plant species are designated by their genus names. Data points are shown as circles; n = 5.

uniform in biomass each species in the community was. It was

calculated using the function “diversity” from the package
“vegan”.33 The larger the differences in biomass between
species the less uniform the community, as some species will
dominate.34 All statistical analyses were done with R version
3.5.3.35 Results shown throughout the text are mean values ±1
standard error (SE).
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■ RESULTS
Plant community biomass was independently affected by
drought and microfibers (Table 1). Community shoot and root
mass decreased from well- watered to drought conditions by
∼47% and ∼53%, respectively, but by constrast, shoot mass
increased by ∼6% and root mass by ∼90% when the soil had
added microfibers (Figure 1a,b). Community root:shoot ratio
increased when microfibers were added to the soil and was not
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Figure 4. Microplastic fibers and drought effects on (A) root diameter (RAD), (B) root tissue density (RTD), (C) specific root length (SRL), and
(D) specific root surface area (SRSA). The box in each boxplot shows the lower, median, and upper quartile values, and the whiskers show the
range of the variation. Data points are shown as circles; n = 5.
significantly affected by drought (Table 1, Figure 1c). Plant
community structure, in terms of biomass, was affected by
drought and microfibers (Figure 2, Table 1, Figures S2−S4).
All plants survived until the end of the experiment. Drought
decreased biomass for all plant species, while microfibers had a
different effect depending on plant species identity and
drought conditions (Figure 3). Microfibers added to the soil
increased shoot mass of Calamagrostis by ∼66% under well-
watered conditions and of Hieracium by ∼85% under drought
conditions, but microfibers decreased shoot mass of Holcus by
∼78% under well-watered conditions and of Festuca by 51%
under drought conditions (Figure 3). Community evenness, in
terms of biomass, was only affected by microfibers (Table 1),
which increased when the microfibers was added to the soil.
Under drought conditions (with and without microfibers),
evenness was similar to that under well-watered conditions
with microfibers (Figure 1d). At the community level, root
traits were affected by drought but not microfibers (Table 1,
Figure 4). Root diameter increased by ∼24% with drought
while RTD decreased by ∼83% (Figure 4a,b). Although no
significant changes were detected for SRL or SRSA (Table 1),
both tended to decrease when microfibers were added to the
soil (Figure 4c,d). SRL and SRSA decreased by ∼88% and
94%, under well-watered conditions, and by 81% and 85%,
under drought conditions, respectively. Soil bulk density
decreased by ∼12% when microfibers were added in the soil
(Figure S3)
■
DISCUSSION
Plant Productivity. Drought negatively affected plant
biomass (a proxy of productivity36), while microfibers in soil
had the opposite effect. Although the threshold of drought
depends on the species identity and its environmental
adaption, the reduced shoot mass due to drought is a well-
known phenomenon that can be generalized across plant
species,37 as photosynthesis is one of the key processes to be
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affected by water deficit. Contrary to the shoot response, the
response of root biomass to drought is less predictable as it
varies depending on the plant species identity (e.g., plant
functional group),17,38 but at the level of plant community, we
observed that root biomass was reduced by drought while it
was increased by microfibers. Although some species tend to
invest more biomass into longer-lasting root organs, in order to
optimize water uptake over time,39 our pattern aligns with the
idea that biomass of fine roots is often reduced with drought as
a consequence of reduced transpiration and respiration
rates.40,41 Additionally, it is likely that the increase in root
biomass in soil with microfibers can be linked to reduced soil
bulk density and an increase in soil macroporosity,42,43 which
ultimately, improved aeration14,16 and facilitated a better
penetration of roots in the soil matrix.44 Root mass increases
will promote water and nutrient uptake, rhizodeposition,
microbial activity, and mycorrhizal associations,20,45,46 which
translate to an increase in shoot biomass. In terms of gauging
effect sizes, plant communities under drought conditions with
microfibers had similar root biomass as those under well-
watered conditions without microfibers. Water availability
would increase with microfibers in the soil, as fibers would
keep water content higher for longer periods, as was observed
in soils similar to ours.14 Soil water holding capacity is strongly
related to soil microporosity and thus to soil bulk density.43
Additionally, as water holding capacity is positively correlated
with soil aggregation,47 and microfibers may contribute to this
soil property by helping to entangle soil particles,16 micro-
plastic fibers would indirectly promote water holding capacity
by their positive effects on soil aggregation. All these potential
positive effects of microfibers on WHC would suggest an
amelioration of drought conditions. However, biomass increase
was only seen for root and not for shoot biomass, which
suggests that microfibers did not sufficiently ameliorate the
negative effects of drought at the level at which we applied
drought here. Along with plant biomass, drought also modifies
root morphological traits.17,48 Our results showed that drought
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but not microfibers affected root morphological traits at the
community level. Root diameter increased while root tissue
density decreased with drought, which can be linked with the
fact that roots with these characteristics may support faster
resource acquisition with a lower investment.49,50 We did not
find a clear response in SRL or SRSA, traits that are typically
highly responsive to drought,17,18 likely due to the presence of
different species that counteract the effect of drought.
However, our data showed that at community level, SRL and
SRSA tended to decrease, likely in order to diminish the risk of
hydraulic rupture and face drought.51,52
Our results showed that in the short term, microfibers
increased plant productivity and modified root morphological
traits; however, we do not currently know what the long-term
responses will be, as additional factors could come into play.
For instance, the increase in root biomass may create channels
in the soil profile, which having been influenced by the drought
conditions could facilitate downward movement of the
microfibers.27,53 Additionally, microplastics, depending of the
degradability, hydrophobicity, electric charge, or roughness,
can also serve as vectors for pathogens, as their surface may
harbor potential pathogenic bacteria.54,55 Thus, when micro-
plastics are added into the soil, pathogens would easily be
translated into the soil matrix, as seemingly they do not
distinguish between natural (e.g., wood) and artificial
surfaces.55 After some time, the accumulation of soil pathogens
due to microplastics can suppress plant growth causing
negative plant−soil feedback.56 In addition, toxic substances
already present or absorbed onto microplastics57,58 would
negatively affect not only plant roots (growth, and thus
rhizodeposition) but also soil biota composition and microbial
activity, which could then secondarily affect plant growth.
Likewise, the high carbon content of microplastic59 could lead
to a microbial N immobilization, with consequences for plant
community productivity and composition.
Plant Community Structure. Plant community structure,
here, in terms of biomass per species in our community
modules, was affected by drought and microfibers in the soil.
Drought stress was such that all plant species had reduced
biomass irrespective of the presence of microfibers in soil,
while the effect of microfibers on plant growth was variable and
depended on the water conditions and plant species. Under
drought conditions and with microfibers in soil, Festuca had
decreased growth, while Hieracium, a species that through
allelopathy can reduce the germination and growth of
neighboring species,60 had increased growth. This suggests
that under drought conditions, the decrease of soil bulk density
due to microfibers led to a better performance of Hieracium,
which may in turn have had a detrimental allelopathic
influence on Festuca.
Microfibers in the soil promoted the growth of the highly
invasive species Calamagrostis, especially under well-watered
conditions. Calamagrostis is a species that is widespread across
the whole of temperate Eurasia. It uses the guerrilla strategy of
clonal growth to spread rapidly, reducing community diversity
by competitive exclusion of other plant species.61 In fact,
Calamagrostis has invaded many high-value seminatural
grasslands of Central Europe, contributing to biodiversity
decline.62 Different efforts have been dedicated to counteract
the growth of this species: mowing, grazing, or biological
control using hemiparasitic plants.62,63 However, efforts to
control this invasive species could be made more difficult if
Calamagrostis in fact grows better with microfibers in the soil,
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whose strategy of clonal growth could promote its dominance
in the field.
Moreover, microfibers tended to decrease biomass of Holcus,
albeit slightly. Holcus is a native species in Europe with a rapid
clonal growth64 that would facilitate the establishment of
different plant species through the amelioration of micro-
climatic conditions65 as it performs well in bare soils and
disturbed areas.64 Likewise, species such as Holcus could
potentially reduce invasiveness of Calamagrostis by creating a
more competitive environment for that species. The negative
effect of microfibers on Holcus in our experiment could thus
have contributed to the increased success of Calamagrostis.
Our results clearly showed contrary responses of plant
species growing in a community to microfibers in the soil.
Future research on this topic should include different plant
species and growth forms (e.g., trees) and explore responses
over more than one growing season. Additionally, future
research should explore the effects of other microplastics on
different plant communities to achieve a greater generalizability
of the patterns observed here.
Potential Consequences for Ecosystem Functioning.
Plant communities often respond more rapidly to human
activities or altered environmental constraints in terms of
evenness than those of richness.66 Our results show that for
our selected species, microfibers added to the soil at 0.4%
concentration, without drought, strongly affect community
evenness, in terms of biomass, which could impact ecosystem
functions. When synergistic interactions are important for the
community, reduced evenness may have negative consequen-
ces,67 but when an ecosystem function is highly dominated by
few species,68 an increase in evenness could reduce ecosystem
functionality. Microplastics can not only modify community
evenness, in terms of biomass, but also potentially affect
community evenness in terms of species abundance or survival.
Litter decomposition, which is a key process enabling the
recycling of carbon and nutrients, will be altered, as litter
production and quality change along with plant relative
abundances.69 Soil properties, such as aggregate stability,
aeration, and moisture content, which are strongly affected by
microplastics,14 would influence the accessibility of soil carbon
to active decomposer enzymes70 and thus litter decomposition
and soil carbon storage. Additionally, in communities that are
dominated by species highly resistant or resilient to
disturbance, a change in relative abundance of plants could
affect community resistance or resilience.36 Such effects on
ecosystem functioning should be a high priority for future
research. In addition, it will be important to test effects of not
only microplastic fibers, which are highly abundant worldwide,
but also other microplastic types that may have potential
consequences for ecosystem functioning; for instance: micro-
plastic films in agricultural lands or microplastic fragments
(e.g., car tires) in urban ecosystems. Future research should
also include different concentrations of microplastics in the soil
as they may vary depending on the proximity to the emitting
source, the microplastic type, and the ecosystem studied.
■
*
ASSOCIATED CONTENT
sı Supporting Information
The Supporting Information is available free of charge at
https://pubs.acs.org/doi/10.1021/acs.est.0c01051.
Microplastic fiber size distribution (Figure S1); photo-
graphs of the experimental setup (Figure S2); micro-
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Environ. Sci. Technol. 2020, 54, 6166−6173
Environmental Science & Technology
plastic fiber effects on bulk density (Figure S3);
properties of polyester fibers (Table S1); results from
linear models on shoot mass per plant species (Table
S2); eigenvalues of the principal component analysis for
plant community structure (Table S3); results from
multivariate analysis of variance of plant species biomass
(Table S4) (PDF)
■
AUTHOR INFORMATION
Corresponding Author
Yudi M. Lozano − Freie Universität Berlin, Institute of Biology,
Plant Ecology, D-14195 Berlin, Germany; Berlin-Brandenburg
Institute of Advanced Biodiversity Research (BBIB), D-14195
Berlin, Germany; orcid.org/0000-0002-0967-8219;
Email: yudyja@gmail.com
Author
Matthias C. Rillig − Freie Universität Berlin, Institute of Biology,
Plant Ecology, D-14195 Berlin, Germany; Berlin-Brandenburg
Institute of Advanced Biodiversity Research (BBIB), D-14195
Berlin, Germany; orcid.org/0000-0003-3541-7853
Complete contact information is available at:
https://pubs.acs.org/10.1021/acs.est.0c01051
Author Contributions
Y.M.L. and M.C.R. conceived the ideas and designed
methodology; Y.M.L. established and maintained the experi-
ment in the greenhouse; Y.M.L. analyzed the data and led the
writing of the manuscript. All authors contributed critically to
the drafts and gave final approval for publication.
Notes
The authors declare no competing financial interest.
■
ACKNOWLEDGMENTS
The work was funded by the German Federal Ministry of
Education and research (BMBF) within the collaborative
Project “Bridging in Biodiversity Science (BIBS-phase 2)”
(funding number 01LC1501A). M.C.R. acknowledges addi-
tional support from an ERC Advanced Grant (694368). We
thank Christine Mhundwa, Tydings Morrison, and Jenny M.
Ospina for their assistance in the experimental setup and data
collection. The experimental design was greatly improved by
discussions with Carlos Aguilar-Trigueros, Gabriela Onaindia,
Carlos Acame, and Stefanie Maaβ. The authors declare no
competing financial interest.
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