Urban Forestry & Urban Greening 41 (2019) 170–178

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Urban Forestry & Urban Greening

journal homepage: www.elsevier.com/locate/ufug

Water relations of street trees in green infrastructure tree trench systems T

⁎
Joshua S. Caplan, Russell C. Galanti, Stuart Olshevski, Sasha W. Eisenman
Temple University, Department of Landscape Architecture & Horticulture, 580 Meetinghouse Road, Ambler, PA, 19002, USA

A R T I C LE I N FO A B S T R A C T

Handling Editor: Justin Morgenroth Urban trees provide a number of ecosystem services through the process of transpiration, including evaporative
Keywords: cooling and returning stormwater to the atmosphere. Collocating street trees and engineered stormwater
Acer × freemanii catchments in ‘tree trenches’ is a space-efficient strategy for providing these services and managing stormwater
Infiltration trench simultaneously; such systems are therefore being installed in a growing number of cities worldwide. However,
Platanus × acerifolia tree trenches are designed to infiltrate rapidly, which could limit water availability in trenches and soil pits,
Stomatal conductance thereby restricting transpiration between storms and eventually impacting tree health and survival. Focusing on
Stormwater control measure tree trenches in Philadelphia, USA, we sought to determine how the water relations of young trees varied with
Urban forestry
atmospheric and soil moisture conditions, if water limitation affected photosynthesis and basal growth, and
whether interspecific differences in water use were related to leaf economics. Measurements of stomatal con-
ductance (gs) from 13 species and cultivars indicated that water use varied widely across taxa; mean gs ranged
from 161 to 507 mmol m−2 s-1, as did the responsiveness of gs to increasing vapor pressure deficit. Mean gs was
correlated with photosynthesis across all taxa, with gas exchange appearing to influence growth rates in the
majority of species. We found no evidence that the sorting of taxa was related to leaf economics, but it was
modestly consistent with prior assessments of tolerance to xylem cavitation risk. Strong differences in leaf water
potential between the two taxa for which it was measured (Acer × freemanii and Platanus × acerifolia, both of
which are isohydric) suggested that some taxa may have avoided water limitation by accessing supplemental
water sources such as the soil beneath trenches. These results demonstrate that young individuals of common
street tree taxa vary considerably in their ability to maintain favorable water relations through hot and dry
periods in tree trenches as they are typically designed.

1. Introduction
A number of cities with combined sewer systems are coupling their
efforts to manage stormwater and to expand urban forests. Combined
sewer systems collect stormwater runoff and sewage in a single network
of pipes, delivering the effluent to treatment plants under most cir-
cumstances (US EPA, 2004). However, when large storms cause the
volume of wastewater to exceed the capacity of treatment plants, ef-
fluent is redirected into nearby waterways; approximately 3.2 trillion
liters of untreated sewage and stormwater are released into the en-
vironment through combined sewer overflow events in the United
States each year (US EPA, 2004). To address this issue, a number of
cities are building distributed networks of green stormwater infra-
structure (GSI) systems to capture stormwater before it enters combined
sewers; common types of GSI systems include bioswales, rain gardens,
green roofs, and tree trenches (Sabbion, 2018). Although designs vary
considerably, most GSI systems hold water in soil- or gravel-filled beds,
allowing it to infiltrate or be transpired by plants growing at the
surface.
While most GSI systems primarily contain shrubs and herbaceous
plants, the installation of tree trenches contributes directly to urban
forest expansion. For example, in the process of installing approxi-
mately 350 tree trenches between 2011 and 2018, Philadelphia, USA
added around 1000 street trees to its urban forest (Philadelphia Water
Department, unpublished data). Aboveground, a tree trench looks like a
typical row of street trees and sidewalk, but belowground it contains an
elongated, gravel-filled catchment (Fig. 1); the trees’ soil pits are em-
bedded in the gravel, though typically separated by a geotextile
(Gaines, 2016; Grohmann and Menconi, 2016). Stormwater runoff from
adjacent roadways and other impervious surfaces enter at one end of
the trench and are distributed through it via a perforated pipe. The
design of tree trenches is intended to allow stormwater in the trench to
reach the tree pits during sufficiently large storms, essentially watering
them from beneath. This is expected to increase the amount of water
available to trees in the hours to days following storms relative to
traditional soil pits (Grohmann and Menconi, 2016; Philadelphia Water

⁎
Corresponding author.
E-mail address: eisenman@temple.edu (S.W. Eisenman).

https://doi.org/10.1016/j.ufug.2019.03.016
Received 20 September 2018; Received in revised form 21 February 2019; Accepted 27 March 2019
Available online 28 March 2019
1618-8667/ © 2019 Elsevier GmbH. All rights reserved.
J.S. Caplan, et al.
Department, 2018). In principle, street trees in these integrated systems
provide the same ecosystem services as trees in other contexts (e.g.,
heat reduction, aesthetic improvement, particulate filtration) but also
enhance stormwater interception, infiltration, and removal (Kuehler
et al., 2017).
Although the potential benefits of collocating street trees and be-
lowground stormwater catchments are clear, there are potential draw-
backs as well. For one, trees may experience more severe water lim-
itation between storms if embedded in GSI trenches. This could occur if,
after the successful infiltration of stormwater, the void spaces in gravel
beds are filled with air for extended periods of time (Grabosky et al.,
2009). In addition to reducing water availability directly, this could
cause soil in tree pits to lose water more rapidly than would occur in
tree pits surrounded by native soil (McCartney et al., 2005). Thus, for
young trees to gain access to a relatively consistent water source at
depth, they must first grow beyond both the soil pit and gravel bed and
establish roots in the soil below the trench (Bartens et al., 2009). In
addition, soils could potentially reach near-saturated conditions im-
mediately following large storms, potentially inducing flooding re-
sponses such as hypoxia and ion toxicity (Willey, 2016). Further, the
ability of trees to maintain active transpiration, and therefore photo-
synthesize and grow, in the unique hydrologic conditions of a street tree
pit depends strongly on atmospheric conditions such as the evaporative
demand of the air, solar irradiance, and wind speed (Moser-Reischl
et al., 2019; Rahman et al., 2017; Whitlow et al., 1992). Characterizing
tree performance in response to varying environmental conditions
could therefore provide a means of assessing the circumstances under
which a typical trench design facilitates or limits water availability for
trees.
Tree performance in GSI trenches can also be expected to vary
across taxa, as adaptations to growth in particular environments may be
conferred from species’ evolutionary histories. For example, a species’
ability to tolerate periods of low water availability could be influenced
by its propensity to continue transpiring when there is an increased risk
of xylem cavitation (i.e., its isohydric vs. anisohydric status; Attia et al.,
2015). In addition, traits associated with resource use may be predictive
of organismal-scale growth rates and whether a species is likely to
proliferate roots into the soil below a trench before water in the soil pit
becomes limiting to further growth. Such traits could include those
reflecting the tradeoff between the energetic cost of leaf tissue con-
struction and leaf longevity (i.e., the leaf economics spectrum; Wright
et al., 2004), such as specific leaf area (SLA; the ratio of leaf area to leaf
dry mass). Given the mounting evidence that, at least in woody plants,
there is a spectrum of ‘fast’ to ‘slow’ strategies for resource acquisition
and use due to similar tradeoffs in stem and fine root construction
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Urban Forestry & Urban Greening 41 (2019) 170–178
Fig. 1. Elements of a green stormwater infrastructure
tree trench (after Tu and Traver, 2018a). The system
depicted was the largest used in this study and the
subject of intensive hydrological and water relations
data collection. The trench is shown in section view
(top) and plan view (bottom). Dashed lines depict the
extent of soil pits below the sidewalk.
(Reich, 2014), it is plausible that easily measured morphological traits
(like SLA or wood density) could be predictive of physiological cap-
abilities (like photosynthetic rates and water use efficiency) and growth
potential. Of course, functional traits are also influenced by evolu-
tionary histories and local conditions, but it is often possible to discern
broad-scale patterns of adaptation from traits (Adler et al., 2014;
Caplan et al., in press).
We sought to determine how tree species and varieties commonly
used in urban street-side settings respond to variation in environmental
conditions when planted in GSI trenches. We specifically focused on
plant water relations (i.e., stomatal conductance and leaf water po-
tential), as these provide a means of assessing the net effects of phy-
siological and morphological adaptations to varying water availability
and atmospheric conditions (Caplan and Yeakley, 2010; Close et al.,
1996; Whitlow et al., 1992). Our first objective was to characterize
interspecific variation in water use for young street trees planted in
stormwater trenches. We did this by determining how stomatal con-
ductance varied across taxa and over a broad range of vapor pressure
deficit (VPD, a measure of the atmosphere’s evaporative demand).
Second, we sought to determine how simultaneously varying environ-
mental conditions (i.e., soil water availability, VPD, wind speed, and
solar radiation) collectively influence street tree water relations in the
context of a GSI trench. To do this we conducted an expanded analysis
with two of the species in a trench whose environmental conditions
were monitored intensively. Our third objective was to determine if
interspecific differences in water use can explain differences in tree-
level growth, and if variation in one or both factors can be predicted by
plant functional traits associated with resource use economics. Speci-
fically, we investigated correlations between characteristics in our gs
dataset (i.e., mean and slope in the VPD relationship) and patterns of
trunk growth, photosynthesis, and specific leaf area.
2. Methods
2.1. Field site
The focus of this study was a set of 26 street trees in Philadelphia,
USA (40.0701 °N, 75.1753 °W). The trees were located in 10 storm-
water trenches along two-lane streets adjacent to a school’s parking lot,
its athletic field, and the surrounding residential area. Trees were
planted in 2014 and had 8.3 ± 1.6 cm diameter trunks (mean ± SD;
measured ˜75 cm above ground level) at the end of the 2015 growing
season. There were 13 species or cultivars from 11 genera represented
in the set (Table 1); two taxa of both Acer (maple) and Quercus (oak)
were included. This set of trees was therefore appropriate for making
J.S. Caplan, et al.
Table 1
Tree taxa (species and cultivars) included in the study together with the number
of individuals evaluated.
Scientific name Common name Number
Acer × freemanii 'Armstrong' Freeman's maple 2 evaporative demand of the atmosphere, we calculated VPD at all time-
Acer rubrum Red maple 2
Amelanchier laevis 'Snowcloud' Snowdrift crabapple 2
Cercis canadensis 'Forest pansy' Eastern redbud 2 turated vapor pressure of water at Tamb (Wagner and Pruss, 1993).
Cornus florida 'Cherokee chief' Dogwood 1 Volumetric soil water content (θv) was also recorded in each of the five
Crataegus viridis Hawthorn 2 soil pits (5 min interval) using Stevens Hydra Probe II soil moisture
Koelreuteria paniculata Golden rain tree 2
Malus 'Sentinel' Sentinel crabapple 2
Platanus × acerifolia 'Bloodgood' London plane tree 3
Prunus sargentii Sargent cherry 2 depth. However, data gaps and uncertain measurement error precluded
Quercus macrocarpa Bur oak 2 the use of individual time series for each tree. Instead, we derived a
Quercus robur English oak 2 single, complete time series by imputing data for one tree at 35 cm
Syringa reticulata 'Ivory silk' Japanese tree lilac 2
interspecific comparisons, though the number of replicate trees per
taxon (n = 1–3) was too small to rule out the possibility that environ-
mental variation would influence our data.
The ten stormwater trenches we investigated had similar structures
(e.g., Fig. 1), as the design objective in all cases was to capture the first
5 cm of rainwater that fell on the adjacent streets and infiltrate it within
72 h (Philadelphia Water Department, 2014). All trenches were located
beneath sidewalks and contained 2–5 tree pits (3.0 × 1.2 × 0.9 m,
L × W × D) depending on their length (range: 32–82 m) and estimated
drainage area (range: 901-2494 m2). Depths at the shallower ends of
trenches ranged from 1.34 to 1.42 m while depths at deeper ends
ranged from 1.78 to 2.61 m. Trenches were filled almost entirely with a
non-compacting stone aggregate adhering to the AASHTO 57 specifi-
cation (approx. 1.5 cm median diameter, 45% porosity), though trench
bottoms were lined with a 15 cm layer of sand (adhering to AASHTO
10; approx. 1 mm median diameter). Tree pits contained a poorly
graded sandy soil (8.6% organic matter) separated from the sur-
rounding gravel by a geotextile. Each trench had an inlet structure at
one end that received water from one or more drains at street-level. A
perforated distribution pipe (20 cm diameter) ran the length of each
trench to facilitate longitudinal water movement.
Five of the trees were the subject of a more detailed data collection
effort that took place simultaneously with the larger study. These trees
were used to determine how variation in soil and atmospheric condi-
tions can jointly influence the water relations of GSI trees (i.e., our
second objective). The trees were located in a single trench (Fig. 1) in
which hydrologic conditions were monitored intensively (described in
detail by Tu and Traver, 2018b). Three of the trees in this trench were
Platanus × acerifolia 'Bloodgood' (London plane tree) while the re-
maining two were Acer × freemanii ‘Armstrong’ (a red maple-silver
maple hybrid). Both are commonly used street trees in the temperate
urban forests of North America and Europe (Dirr, 2009; Sæbø et al.,
2005) and initial observations suggested that they would exhibit di-
verging water use strategies. The five trees were arranged with the
cultivars in alternating pits along the length of the trench. The trench
itself drained an area of approximately 2494 m2, 88% of which was
paved; it was 82 m long, 2.6 m wide, and its depth varied from 1.42 m
near the inlet to 2.25 m at the opposite end. The variation in depth was
due to the slope of overlying ground surface (1%); the bottom of the
trench had a constant elevation. The distribution pipe (20 cm diameter)
was approximately 1.0 m deep near the inlet and 1.5 m deep at the
opposite end (0.5% slope). Due to the elevation change of the top of the
trench, the distance from the bottom of the tree pits to the trench
bottom varied from 1.5 to 0.7 m.
2.2. Environmental data
A weather station recorded meteorological data at the site at 5 min
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Urban Forestry & Urban Greening 41 (2019) 170–178
intervals for the duration of the 2015 growing season; it was located
2 m from the central tree in the intensively monitored trench.
Monitored variables included air temperature (Tamb), relative humidity
(Rh), wind speed, and total solar radiation (direct plus diffuse; via a
LI200X-L sensor, LI-COR Biosciences, Lincoln, USA). To quantify the
points from Tamb and Rh: VPD = VPsat(1 - Rh), where VPsat is the sa-
sensors (Fondriest Environmental, Fairborn, USA). Sensors were in-
stalled approximately 50 cm from the base of each tree at 10 and 35 cm
depth from the set of partial time series. Specifically, we ran 10 itera-
tions of the predictive mean matching algorithm from the mice code
library (van Buuren and Groothuis-Oudshoorn, 2011) for R 3.4.0 (R
Foundation for Statistical Computing, Vienna, Austria); the mean of the
10 resulting time series (θmn) was used for subsequent analyses. Given
that differences in measured θv among tree pits were small compared to
the range observed during storms, the final time series can be assumed
to accurately represent relative water availability, though values may
be systematically high or low. Water depth in the bottom of the trench
was also measured at 5 min intervals using a HOBO U20-001-04 pres-
sure transducer (Onset, Bourne, USA) placed in an observation well
near the central tree.
2.3. Water relations
Stomatal conductance (gs) was measured in each of the 26 trees
approximately weekly through the 2015 growing season (14 May
through 6 November). More frequent measurements (approximately
twice weekly) were made on the five trees in the intensively monitored
trench; the number of sampling dates ranged from 19 to 41 for most
trees (median = 28) but was 68 or 69 for the five focal trees (n = 891
total tree visits). Data were typically collected between 1100 and
1300 h (always between 1030 and 1400 h) and only on days when
leaves were dry. A single SC-1 Leaf Porometer (Meter Environment,
Pullman, USA) was used for all measurements and was calibrated daily.
During tree visits, gs measurements were made on three leaves from the
portion of each tree’s mid-canopy receiving the most sunlight; means of
the triplicate values were used for further analysis.
Mid-day leaf water potential (Ψmd) was measured on the five focal
trees on all measurement days (n = 69 per tree). Measurements were
made within ˜5 min of corresponding gs measurements; they therefore
also occurred during mid-day or early afternoon (typically
1100–1300 h). One to two fully exposed leaves from the middle part of
each tree’s canopy were excised per tree and immediately inserted into
a Model 615 Pressure Chamber (PMS Instruments, Albany, USA) for
Ψmd measurement (Gonzáles, 2001).
Prior to conducting statistical evaluations we extracted environ-
mental data (e.g., VPD, θmn, wind speed, and solar irradiance values)
that were collected nearest in time to water relations measurements.
Tree-level means were then calculated for data collected on the same
day.
2.4. Growth and trait data
Tree growth rates were determined from increases in trunk diameter
over a two year period. Using a caliper or DBH tape, trunk diameters
were measured approximately 75 cm above ground level on a single day
in the winters following the 2015 and 2017 growing seasons. The dif-
ference in natural log-transformed values was calculated as a metric of
relative growth.
Leaf trait data came from a combination of records from the TRY
J.S. Caplan, et al.
plant trait database (Kattge et al., 2011) and measurements in the field.
Data describing photosynthetic rates under light-saturated conditions
(Asat) and SLA were publicly available through TRY for 7 and 9 of the
focal species or closely related taxa, respectively, with the number of
records pertaining to a species varying from 1 to 210 (Table S1). We
also sought data on traits related to wood anatomy, but there was in-
sufficient coverage at the species and record level to justify proceeding.
To fill gaps in the Asat and SLA datasets and to ensure that trees from the
site were represented, we measured Asat on each of the 26 trees on a
single day in July 2018 and collected leaves for SLA determination
concurrently. Asat was measured using an LI-6800 photosynthesis
system (LI-COR Biosciences, Lincoln, USA) that was not previously
available. For these measurements, leaf temperature was held at 30 °C,
irradiance at 2000 μmol m−² s−¹, reference [CO2] at 400 ppm, and Rh
between 60 and 75%. For SLA, fully expanded leaves (n = 9–11) were
collected and later measured for total leaf area (with an LI-3000, LI-
COR Biosciences, Lincoln, USA) and total dry mass; SLA was calculated
as the ratio of total area to total mass. TRY and field data were com-
bined by computing means across all available data; the mean value
from the trees in the field and individual records from TRY were given
equal weight when averaging.
2.5. Data analysis
For comparisons of species-level gs rates and the slopes of species’ gs
vs. VPD relationships (Δgs), daily data were entered into a single mixed-
effects linear model (via function lmer from the R library lme4; Bates
et al., 2015). The response variable was gs, while species, VPD, and
their interaction were explanatory variables. A random effect was also
included to account for repeated measurements within individual trees.
Species’ mean gs rates and the corresponding Δgs values were computed
from this model by estimating marginal means (via the R library em-
means; Lenth, 2017) from each species’ gs data (1) independently of
VPD and (2) while accounting for the gs × VPD interaction, respec-
tively.
A pair of linear mixed-effects models was used to evaluate the
combined effects of belowground water availability and atmospheric
conditions on water relations of trees in the intensively monitored
trench. Response variables were gs or Ψmd, while explanatory variables
included θmn, VPD, solar radiation level, wind speed, species identity,
and all possible interaction effects. A random effect for tree was also
included in both models. Initial models included the depth of water in
the trench, but this led to collinearity among explanatory variables
(variance inflation factors reached 137); water depth was therefore not
included in final models. The significance of fixed effects was evaluated
using Wald χ2 statistics derived from Type II sums of squares (via
function Anova from the R library car; Fox and Weisberg, 2011); all
models used restricted maximum likelihood for parameter estimation.
The relationship between water use and growth was evaluated by
regressing species-level means for basal growth against those of gs or
Δgs. An additional set of models was evaluated that also contained a
two-level categorical variable distinguishing species with normal vs.
high growth for a given gs; this grouping was apparent in visualizations
of the data but not expected a priori. Similarly, relationships between
leaf traits (Asat and SLA) vs. water use parameters (gs and Δgs) were
evaluated via simple regression using species-level means. F statistics
were used to determine if regression coefficients differed from zero.
3. Results
3.1. Variation in water use strategies
Mean ( ± SE) mid-day gs rates varied considerably among species
(χ2 = 244.8, P < 0.001), ranging from 161 ± 22 to 507 ± 22 mmol
m−2 s-1 across the 13 species included in the study (Fig. 2A). The low
and high endpoints of this range were held by Syringa reticulata ‘Ivory
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Urban Forestry & Urban Greening 41 (2019) 170–178
silk’ and Quercus robur, respectively. Pooled across species, gs decreased
with increasing VPD to a modest extent (χ2 = 7.8, P = 0.005), but the
slope of this relationship (i.e., Δgs) was, in fact, highly species-depen-
dent (χ2 = 95.0, P < 0.001; Fig. S1). As inferred from estimates of Δgs
and their 95% CIs, gs declined unequivocally with increasing VPD in
two species: Acer rubrum and Crataegus viridis (Fig. 2B). gs changed little
with varying VPD in eight additional species (i.e., 95% CIs for Δgs in-
cluded zero). Of these, estimates of Δgs were negative in six cases but
positive in two. In the remaining three species, 95% CIs for Δgs ex-
cluded zero but gs increased with increasing VPD; these were Q. robur,
P. × acerifolia ‘Bloodgood’, and C. canadensis 'Forest Pansy'. Together,
the fixed effects in the model explained 44% of the variability in gs,
with the random effect of tree identity raising this to 47%.
The more detailed analysis of P. × acerifolia and A. × freemanii
showed that environmental conditions had strongly interacting effects
on water relations (Fig. 3; Table S2). Ψmd declined as VPD rose
(χ2 = 86.7, P < 0.001) or soil water became less available (χ2 = 46.2,
P < 0.001) in both species, and these effects reinforced one another
(χ2 = 9.3, P < 0.001). Higher wind speeds also exacerbated the effect
of low soil moisture on Ψmd (χ2 = 6.4, P = 0.011), though there was a
non-significant trend indicating that this effect may be stronger for A.
× freemanii than for P. × acerifolia (χ2 = 3.2, P = 0.073). P. ×
acerifolia had a more favorable water status than A. × freemanii on
average (mean ± SE: -1.06 ± 0.018 vs. -1.43 ± 0.035 MPa;
χ2 = 122.1, P < 0.001) and its Ψmd varied less strongly with changes
in either soil moisture (χ2 = 32.1, P < 0.001) or VPD (χ2 = 4.1, P =
0.040). Fixed effects explained 67% of the variation in Ψmd, with
random effects explaining an additional 1%.
The two species had sharply contrasting gs responses to environ-
mental conditions (Fig. 3; Table S2). As indicated above, mean gs was
greater in P. × acerifolia than in A. × freemanii (χ2 = 18.1, P <
0.001) and decreased with increasing VPD for A. × freemanii but in-
creased with increasing VPD for P. × acerifolia (χ2 = 11.2, P <
0.001). In A. × freemanii, gs decreased as soil moisture became less
available (χ2 = 6.5, P = 0.011), much as Ψmd did. In contrast, gs was
minimally affected by soil moisture in P. × acerifolia, at least at the
depth measured. However, the influence of soil moisture on gs de-
pended on VPD (χ2 = 8.2, P = 0.004), with low soil moisture de-
creasing gs in both species more strongly at higher VPDs. In the case of
A. × freemanii, this effect was intensified under higher irradiance
(χ2 = 4.3, P = 0.039). Moreover, both high irradiance (χ2 = 11.1,
P < 0.001) and high wind speed (χ2 = 5.2, P = 0.023) exacerbated
the effect of low soil moisture on gs, though the effect of wind speed was
stronger for A. × freemanii than for P. × acerifolia. Fixed effects ex-
plained 39% of the variation in gs, with random effects explaining an
additional 2%.
3.2. Associations between water use and growth
Species-level growth rates were greater in species that had higher
mean gs (Fig. 4A). While these relationships were statistically sig-
nificant with data from the full set of species pooled (F1,11 = 6.01, P =
0.032, R2 = 0.35), they were substantially improved by the addition of
the categorical variable differentiating species with normal vs. higher
growth at a given gs (F2,10 = 132.8, P < 0.001, R2 = 0.96). Results
were nearly identical when species’ maximum gs rates were used in-
stead of means (data not shown), though this could be expected given
that mean and max gs were highly correlated (r = 0.96). There was no
discernable relationship between tree-level growth and Δgs in the ab-
sence of the grouping variable (F1,11 = 1.21, P = 0.295, R2 = 0.10).
However, with the group term included, there was a clear relationship
wherein the majority of species exhibited a linear increase in growth
rate with increasing Δgs, while the remaining species exhibited near
constant growth rates over a wide range in Δgs (Fig. 4B; F2,10 = 37.8,
P < 0.001, R2 = 0.88).
Interspecific variation in gs had a positive linear relationship with
J.S. Caplan, et al.
Fig. 2. Mean ( ± SE) rates of (A) stomatal conductance (gs) and (B) the slope of the relationship between vapor pressure deficit and gs (Δgs) for each of the tree taxa in
the study. Means are statistically different among pairs of means if bar annotations do not have any letters in common.
variation in photosynthetic rates (Fig. 4C; F1,11 = 8.85, P = 0.013,
R2 = 0.45); the same was true for Δgs but to a lesser degree (Fig. 4D;
F1,11 = 5.78, P = 0.035, R2 = 0.34). There was no discernable re-
lationship between SLA and either gs (Fig. 4E; F1,11 < 0.01, P = 0.925,
R2 < 0.01) or Δgs (Fig. 4F; F1,11 = 0.16, P = 0.693, R2 = 0.02).
4. Discussion
4.1. Variation in water use
Young trees in GSI trenches displayed considerable interspecific
variation in mean gs. Similar ranges of gs have been reported for woody
plants in other studies (Klein, 2014; Poorter and Bongers, 2006; Yi
et al., 2017), though species with substantially faster rates have occa-
sionally been identified (e.g., 800 mmol m−2 s-1 in an invasive shrub;
Caplan and Yeakley, 2010). The lowest mean gs rate observed here
(180 mmol m−2 s-1 for S. reticulata ‘Ivory silk’) is above those typically
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Urban Forestry & Urban Greening 41 (2019) 170–178
Fig. 3. Daily mean values of (A, B) leaf
water potential (Ψmd) and (C, D) sto-
matal conductance (gs) for trees in the
intensively-monitored trench. Means
were calculated from replicate mea-
surements per tree such that a point
represents an individual tree on a given
sampling date. Acer = A. × freemanii
‘Armstrong’ and Platanus = P. × acer-
ifolia ‘Bloodgood’.
measured in plants experiencing moderate drought (< 150 mmol m−2
s-1; Flexas and Medrano, 2002), indicating that these young trees were
not severely water-limited for the majority of the growing season.
However, daily gs rates were indicative of water stress for at least a
subset of measurement days in all species, indicating that soil water was
sometimes insufficient to meet demand (Fig. S1). In addition, the de-
gree to which trees restricted stomatal conductance as VPD increased
(Δgs) varied modestly (remaining between -100 and 100 mmol m−2 s-1
kPa-1), which is also typical of temperate woody species (Franks and
Farquhar, 1999; Waring and Franklin, 1979). Moreover, the 13 species’
gs and Δgs rates were relatively evenly distributed across the range of
values. This may indicate that many manifestations of anatomical and
physiological traits underlie interspecific differences in water relations
rather than one or a few dominant combinations.
Some of the variation in gs and Δgs may be attributable to species
tolerating varying degrees of xylem cavitation risk in the process of
optimizing gas exchange. In evolutionary terms, this has led to distinct
J.S. Caplan, et al. Urban Forestry & Urban Greening 41 (2019) 170–178

Fig. 4. Relationships between water use parameters and those related to growth and leaf economics. Points represent species-level means ( ± SE). Squares denote
species that exhibited relatively rapid rates of basal growth given their mean gs rate, while circles denote species that exhibited normal growth given their mean gs.
Colors used in the online version of this article correspond to those in Fig. 2. Regression lines are shown if linear models indicated that relationships were statistically
significant. SLA = specific leaf area.

water use strategies that can be characterized as isohydric at one ex-
treme and anisohydric at the other (Attia et al., 2015; Sade et al., 2012).
Isohydric plants maintain relatively constant mid-day leaf water po-
tential (Ψlf) and relative water content regardless of water availability
and demand, i.e., they are averse to cavitation risk and therefore re-
strict gs more strongly under drier conditions (typically yielding modest
gs and greater magnitude but negative Δgs). In contrast, anisohydric
plants allow leaf water potential to vary so as to maintain open stomata
across a wide VPD range (typically yielding large gs but Δgs close to
zero); this translates into greater carbon gains but also comes with a
greater risk of cavitation at high VPD or low soil water contents (e.g.,
Moser-Reischl et al., 2019). Of the taxa included here, prior studies
have identified most as isohydric (e.g., Acer spp., Malus spp., Platanus ×
acerifolia), though a few (Quercus spp. and Cornus florida) are known to
be anisohydric (Domec et al., 2017; Jones and Tardieu, 1998; Kjelgren
et al., 2016; Thomsen et al., 2013). The preponderance of isohydric
species may explain why most Δgs values were modest and why Q. robur
and Q. macrocarpa had among the greatest gs rates. Moreover, although
one isohydric taxon, Malus ‘Sentinel’, likewise exhibited relatively rapid
gs, several Malus cultivars are known to exhibit the anisohydric char-
acteristic of delaying stomatal closure until Ψlf is unusually low
(Beikircher et al., 2013). While hydraulic risk tolerance had some

175
J.S. Caplan, et al.
ability to explain variation in gs rates and its response to VPD, other
factors certainly influenced water relations in these GSI trees. This was
clear given that anisohydric C. florida had among the lowest gs rates and
that other taxa in which we measured relatively high gs are isohydric
(e.g., P. × acerifolia).
Another key factor contributing to differences in gs and Δgs may
have been species’ abilities to access water beyond soil pits. It was
particularly striking that P. × acerifolia and A. × freemanii are both
well-known to be isohydric but that only A. × freemanii exhibited ap-
preciable water stress (Ψmd < −2 MPa) when moisture levels in the soil
pits were particularly low (Fig. 3A,B). A plausible explanation for this
contrast in behavior is that A. × freemanii proliferated few roots outside
of the soil pits whereas P. × acerifolia roots extended into the sandy
layer at the bottom of the trench or the native soil below, thus gaining
access to a more temporally stable water source. Consistent with this
possibility, roots were visible in some images of distribution pipe in-
teriors at the time of the study (Tu and Traver, 2018a), suggesting that
this species had proliferated roots well below the soil pits. Our results
therefore support the possibility that low water availability in soil pits
during rain-free periods can be physiologically limiting to trees in GSI
trenches, at least those trenches with relatively rapid infiltration rates
and deep catchments (e.g., compared to the water volume delivered
during a 1 yr storm). Given that soil pit volumes are fixed, the steeply
increasing water requirements that trees have as they grow (Meinzer
et al., 2005) implies that their reliance on supplemental water will ei-
ther have to increase proportionally or growth will have to be re-
stricted. Therefore, the long-term growth trajectory of a tree in a GSI
tree trench may be heavily influenced by both biological and design
factors that determine how readily the trench bottom and native soil
can be accessed. With respect to biological factors, a lack of access to
supplemental water is likely to restrict growth most dramatically in
slow growing species and those that invest minimally in root biomass
when they are young (Caplan and Yeakley, 2013; Drunasky and Struve,
2005; Pallardy and Rhoads, 1993). With respect to trench design,
barriers to growth may be greatest for deep trenches that drain rapidly,
much like the ones studied here.
That several species increased gs with greater VPD levels was sur-
prising, as plants typically maintain constant or decreasing gs with in-
creasing VPD in order to control water loss. This phenomenon may be
attributable to variation in temperature, rather than humidity, dom-
inating the variation in VPD, given that gs is known to increase as
temperatures warm (Urban et al., 2017). The reasons plants would raise
gs in response to warming temperatures are not well established but
thought to include increased evaporative cooling and decreased sto-
matal limitation to photosynthesis (Urban et al., 2017).
Our investigation of gs responses to varying environmental condi-
tions identified multiple interaction effects (Table S2), underscoring the
context-specific nature of plant physiological performance in urban
settings (Moser-Reischl et al., 2019; Salisbury et al., 2018; Whitlow
et al., 1992). In particular, the interaction effects between soil moisture
and other environmental factors indicates that both P. × acerifolia and
A. × freemanii were able to increase gs under high irradiance and VPD
conditions only when soil moisture was sufficient. This further under-
scores the importance of establishing hydrologic connectivity between
the trench and soil pits, which our data suggest has potential for im-
provement. Moreover, our findings reiterate the fact that ratings of
species’ water use capabilities, even those derived from repeated
measurements of physiological rates as we provide here, can only
broadly characterize the degree to which taxa are capable of storm-
water withdrawal. In selecting tree taxa for a given site, consideration
should be given to the size and water retention capacity of the soil pit as
well as the trench depth and infiltration rate (or predictors of it, like soil
porosity), such that species are selected that are well-matched to both
the central tendency and extremes of the prevailing hydraulic regime.
176
Urban Forestry & Urban Greening 41 (2019) 170–178
4.2. Associations between water use and growth
Our analysis indicated that water use was associated with basal
growth rate for the majority of species. However, it also identified a
second group of fast-growing species whose mean gs rates had little to
no relationship with growth. Our results therefore suggest that it may
be possible to predict tree-level growth from gs for species in the slower-
growing group, but it will also be necessary to predict group mem-
bership. This requires first identifying the mechanisms that decoupled
growth from gs, for which we suggest there are three likely possibilities.
First, differences in phenology (i.e., earlier bud break and/or delayed
senescence) may have led to greater annual carbon gains by species in
the more rapidly growing group (Caplan et al., 2015); simple ob-
servations of phenology could be enough to evaluate this possibility,
though analysis of carbon relations may also be needed. Alternatively,
taxa may have had differing access to water, with the more rapidly
growing group having access to plentiful water but the other species
limited by water availability (Pallardy and Rhoads, 1993; Rahman
et al., 2019). Isotopic comparisons of potential water sources and that
in transpiration streams could potentially be used to determine the
relative importance of surficial vs. deep water for across taxa in GSI
trenches (Bertrand et al., 2014). Finally, differences in organismal-level
water use efficiency could have arisen due to differences in boundary
layer conductance, wood anatomy, or other species-specific traits (Bush
et al., 2008; Moser-Reischl et al., 2019). Further evaluation of func-
tional traits in street tree taxa would be valuable in assessing this
possibility, as we found the currently available datasets to be limited in
their taxonomic coverage.
The correlation we observed between gs and Asat confirms that
water use and carbon gain varied in tandem with one another, and
supports the idea that species fall along a spectrum of performance.
However, the lack of a corresponding pattern in SLA makes it difficult
to attribute this variation to leaf economics. It is therefore possible that
environmental factors (like water availability), other factors intrinsic to
the species themselves (like boundary layer conductance, hydraulic
properties of wood, or maximum carboxylation rates), or both, had
substantial influence on gas exchange rates. However, the absence of a
signal corresponding to the fast vs. slow growth grouping in the re-
lationships between gs and both Asat and SLA does suggest that neither
leaf photosynthetic capacity or longevity (a correlate of SLA) were
important in differentiating the two growth groups.
4.3. Design implications
A number of trees in the GSI trenches we studied showed signs of
substantial water stress during hot and dry periods between storms, i.e.,
on days with low soil moisture and high VPD. Specifically, many of the
tree taxa in this study experienced strong reductions in stomatal con-
ductance and, in the case of A. × freemanii and probably others, leaf
water potential declining to levels where the implications can be severe
(e.g., loss of xylem integrity, greater susceptibility to disease, and
stunted growth). We suspect that key factors enabling some taxa to
minimize the effects of water limitation included an ability to pro-
liferate roots into the more temporally stable water in soil beneath
trenches and a greater tolerance of xylem cavitation risk (i.e., aniso-
hydry), possibly coupled with more effective cavitation recovery me-
chanisms. In cases where belowground conditions are similar to those
at our site (e.g., those with deep and rapidly-draining trenches), se-
lecting species with traits that help to avoid or tolerate water stress
(such as P. × acerifolia or Q. macrocarpa) may reduce the ill effects of
water shortage between storms. In principle, tree selection tailored to
the extremes of the hydrologic regime in more mesic or wet GSI tren-
ches would likewise be advantageous.
It may also be possible to modify aspects of typical GSI trench de-
signs such that they enable a greater variety of tree taxa to maintain a
favorable water status during periods between storms. One possibility
J.S. Caplan, et al.
would be increasing infiltration into soil pits from above, potentially by
replacing the overlying concrete sidewalks with permeable pavement or
short-stature vegetation (e.g., grasses), or by directing stormwater from
impermeable surfaces surrounding soil pits towards exposed soil sur-
faces. Increasing the water holding capacity of the soil mix (e.g., by
increasing the clay fraction) may also prolong the period that water is
readily available. Another approach would be to increase the frequency
that soil pits come in contact with stormwater from below, i.e., redu-
cing the water depths required for stormwater to reach soil pits. This
could be accomplished by making trenches shallower or by making soil
pits deeper, though providing soil-filled channels through the gravel
bed may also be feasible in some cases. In addition to providing more
water to soil pits, these adjustments would promote deeper root growth
within soil pits and enable roots to more easily reach native soil.
Additional benefits may come from encouraging downward root
growth, such as fewer conflicts with infrastructure, greater tree stabi-
lity, and improved soil quality at depth.
It is important to note that the combination of rapid drainage rates
and deep gravel beds that likely exacerbated water shortages at our site
simultaneously contributed to the trenches primary functions of storing
and infiltrating stormwater. Efforts to improve tree water relations
through trench design are therefore subject to a tradeoff that may be
best dealt with by maximizing the data available. In particular, in-
formation on sites’ infiltration capabilities under saturated and un-
saturated conditions could be highly informative, especially if used in
hydrologic models to predict the soil water regime as a function of
possible trench depths. This may not yet be possible in the majority of
cases, but could become a realistic practice in the future. In a trench
whose design is fixed, a thorough evaluation of the likely water regime
(e.g., via infiltration measurements before the trench is filled) could
provide valuable information for the tree selection phase. In all like-
lihood, all efforts to optimize trees’ water relations will lead to im-
proved tree health and, therefore, their ability to provide ecosystem
services.
Competing interests
The authors have no conflicts of interest pertaining to the research
described here.
Acknowledgements
This research was made possible by funding from the US
Environmental Protection Agency (grant R835556) and extensive co-
operative support from the Philadelphia Water Department. We are also
grateful for data and input from Peter Tu and other colleagues from the
Villanova Urban Stormwater Partnership. We thank Alyssa Chattin,
Wiley Kollar, and Michael Cappon for assistance with data collection, as
well as Erin Ramsden for contributing database expertise.
Appendix A. Supplementary data
Supplementary material related to this article can be found, in the
online version, at doi:https://doi.org/10.1016/j.ufug.2019.03.016.
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