Science of the Total Environment 442 (2013) 152164

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Science of the Total Environment

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Algal greening and the conservation of stone heritage structures

Nick A. Cutler a, b,, Heather A. Viles a, Samin Ahmad a, Stephen McCabe c, Bernard J. Smith c
a
School of Geography and the Environment, Oxford University Centre for the Environment, South Parks Road, Oxford, OX1 3QY, UK
b
Department of Geography, University of Cambridge, Downing Place, Cambridge, CB2 3EN, UK
c
School of Geography, Archaeology and Palaeoecology, Queen's University Belfast, Elmwood Avenue, Belfast BT7 1NN, UK

H I G H L I G H T S

We investigated green algal biolms on four sandstone heritage structures in Belfast.

The relationships among greening and environmental variables were complex.
Green biolms had little or no impact on the physical integrity of building stone.
The inuence of green biolms on moisture levels in stone may be bioprotective.

a r t i c l e i n f o a b s t r a c t

Article history: In humid, temperate climates, green algae can make a signicant contribution to the deterioration of building
Received 10 August 2012 stone, both through unsightly staining (greening) and, possibly, physical and chemical transformations.
Received in revised form 10 October 2012 However, very little is known about the factors that inuence the deteriorative impact and spatial distribu-
Accepted 11 October 2012
tion of green algal biolms, hindering attempts to model the inuence of climate change on building conser-
Available online xxxx
vation. To address this problem, we surveyed four sandstone heritage structures in Belfast, UK. Our research
Keywords:
had two aims: 1) to investigate the relationships between greening and the deterioration of stone structures
Green algae and 2) to assess the impacts of environmental factors on the distribution of green biolms. We applied an
Biodeterioration array of analytical techniques to measure stone properties indicative of deterioration status (hardness, colour
Weathering and permeability) and environmental conditions related to algal growth (surface and sub-surface moisture,
Algal soiling temperature and surface texture). Our results indicated that stone hardness was highly variable but only
Building stone weakly related to levels of greening. Stone that had been exposed for many years was, on average, darker
Climate change and greener than new stone of the same type, but there was no correlation between greening and darkening.
Stone permeability was higher on old, weathered stone but not consistently related to the incidence of
greening. However, there was evidence to suggest that thick algal biolms were capable of reducing the in-
gress of moisture. Greening was negatively correlated with point measurements of surface temperature, but
not moisture or surface texture. Our ndings suggested that greening had little impact on the physical integ-
rity of stone; indeed the inuence of algae on moisture regimes in stone may have a broadly bioprotective
action. Furthermore, the relationship between moisture levels and greening is not straightforward and is
likely to be heavily dependent upon temporal patterns in moisture regimes and other, unmeasured, factors
such as nutrient supply.
2012 Elsevier B.V. All rights reserved.

1. Introduction known to contribute to the degradation of stone heritage structures

A substantial proportion of the world's tangible heritage is
constructed from stone (Scheerer et al., 2009): it is essential to under-
stand the biogeochemical processes that occur on and in stone struc-
tures if this heritage is to be effectively conserved. Lithobiotic
(stone-dwelling) microorganisms are key players in many of the bio-
geochemical processes that occur in stone. Bacteria and fungi are
Corresponding author at: Churchill College, Cambridge, CB3 0DS, UK. Tel.: +44 1223
336202; fax: +44 1223 336180.
E-mail address: nac37@cam.ac.uk (N.A. Cutler).
(referred to as stone hereafter) through both biophysical and bio-
chemical means (Gaylarde et al., 2003; May et al., 1993; McNamara
and Mitchell, 2005; Warscheid and Braams, 2000). However, in cer-
tain areas (notably those with cool, humid climates) sub-aerial
green algal species may also make a major contribution to the deteri-
oration of stone (Gaylarde and Gaylarde, 2005). Green staining
(greening) resulting from the chlorophyll in green algal cells is a
common feature on stone structures in temperate latitudes (in the
Tropics, cyanobacteria may also make a major contribution to greening:
see Gaylarde and Gaylarde, 2005) and there is also evidence that
sub-aerial green algae are connected with the physical disintegration

0048-9697/$ see front matter 2012 Elsevier B.V. All rights reserved.
http://dx.doi.org/10.1016/j.scitotenv.2012.10.050
 N.A. Cutler et al. / Science of the Total Environment 442 (2013) 152164 153

of stone. However, little is known about the overall contribution that
green algae make to the biodeterioration of stone and the ecological
factors that inuence their growth on stone surfaces (Cutler and
Viles, 2010). This undermines efforts to model the role of algal biode-
terioration under altered climatic regimes. Based on this, the overall
objectives of this research were to 1) investigate the relationships
among green algal growth and the deterioration of building stone,
including impacts on the movement of moisture and 2) to investi-
gate the relationship among environmental factors and the distribu-
tion of green biolms on stone buildings (Table 1). To achieve these
aims, we surveyed four stone heritage structures in Belfast, UK, and
compared the properties of stone surfaces with and without algal
cover.
The biodeterioration of stone has both aesthetic and physical com-
ponents. Changes in surface appearance related to organic activity are
described as biofouling; biological activity that directly contributes to
the physical breakdown of the stone is termed bioweathering (Cutler
and Viles, 2010). The aesthetic impact of green algal biolms is obvi-
ous and well understood. In addition to staining due to pigments
present in algal cells, the formation of green biolms also enhances
the entrapment of particulates which darken the stone surface over
time (Viles and Gorbushina, 2003; Viles et al., 2002). Both the partic-
ulates and carbon xed by the algae provide a source of nutrition for
heterotrophic microorganisms which may further degrade the stone
surface (Gorbushina et al., 1993; Saiz-Jimenez, 1997).
In contrast to the readily apparent impact of green biolms on the
appearance of stone, the physical and chemical impact of green algae
on stone is largely unknown. There have been reports linking green
algae to biophysical deterioration of stone. Wakeeld et al. (1996),
for example, observed granular disintegration of a sandstone struc-
ture associated with growth of ne, lamentary structures of algae
from the genus Trentepohlia. The expansion and contraction of algal
cells associated with wetting/drying or freeze/thaw cycles may also
contribute to the weathering of stone substrates (Hall and Otte,
1990). Many microorganisms, including bacteria and fungi (both
free-living and as lichen symbionts) contribute to stone weathering
by biochemical means, e.g. the production of organic acids (Bock
and Sand, 1993; McNamara and Mitchell, 2005). Green algae are
known to produce small quantities of organic acids (John, 1988)
and may contribute to biochemical degradation of stone by increasing
concentrations of dissolved CO2 through respiration (Welton et al.,
2003). As well as direct impacts on structure, green algal biolms may
also have indirect effects on the stone substrate, most notably on the in-
gress and egress of moisture. Sub-aerial green algae are highly effective
at sequestering moisture; algal cells expand when hydrated and turgid
algal cells could block pore spaces, forming a surface layer of low per-
meability. If this process occurs, a green algal biolm would retain mois-
ture at the stone surface and prevent the egress of deep-seated
moisture, meaning that both the surface and sub-surface of the stone
would remain wetter for longer periods. Water is key component in
stone weathering: it is therefore possible that green algal biolms
have negative repercussions for both the appearance and long-term sta-
bility of stone heritage structures.
In order to model the impacts of green biolms, it is necessary to
understand the ecology of green algae living on stone surfaces, at
both macro- and microscales. On a large scale, sub-aerial green
algae have been demonstrated to respond to uctuations in climate.
Environmental change during the twenty rst century is therefore
likely to alter the relative importance of green algae as agents of bio-
deterioration (Viles and Cutler, 2012). Changes in moisture supply
appear to be particularly signicant (Bellinzoni et al., 2003). Algal bio-
deterioration is likely to become more prevalent in temperate areas
receiving increased rainfall, particularly if a greater proportion of
the rainfall occurs in extreme events. Under this scenario, stone struc-
tures are likely to be subject to prolonged wetting and, presumably,
dramatic increases in the density and extent of green algal biolms.
Anecdotal evidence suggests that this process may be already under-
way in parts of the NW UK (Smith et al., 2010).
Algal greening is also sensitive to microenvironmental variation. Sur-
face texture has been linked to biolm formation in previous studies
(Barberousse et al., 2006; Tomaselli et al., 2000) and is likely to inuence
bioreceptivity (the suitability of a surface for biological colonisation).
Surface irregularities inuence surface drainage and may provide
microsites for algal colonisation i.e. rough surfaces are likely to have
higher bioreceptivity than smooth substrates (Guillitte, 1995). Similarly,
anecdotal evidence suggests that green biolms are more prevalent
close to ground level, where moisture is available from rain-splash and
rising dampness.
1.1. Study area
We conducted our study in central Belfast, UK. Belfast has a large
stock of sandstone heritage structures, mostly dating from the mid-
to late-nineteenth century (Curran et al., 2010). Sandstone (Sst) is
the dominant building stone in NW UK and is globally signicant as
a material used in the construction of heritage structures (UNESCO,
2011). Belfast has a temperate, maritime climate with mild winters
and cool summers. Mean annual minimum and maximum tempera-
tures range between 5.8 and 12.5 C; average annual precipitation is
around 860 mm and there are ~ 155 days with precipitation each

Table 1
The objectives of the study, along with allied hypotheses and analytical approaches.

Hypotheses Analysis

Objective 1: greening and biodeterioration

Physical integrity: Algal greening is likely to promote the physical
disintegration of stone, through both biophysical and
biochemical processes.
Appearance: Green biolms are likely to promote the darkening of
stone surfaces, by trapping particulates and facilitating the
development of other microbial communities.
Movement of moisture: Green algal cells swell when wet, blocking
pore spaces, lowering surface permeability and trapping
sub-surface moisture.
H1: The degree of greening will be positively correlated
with variability in stone surface hardness.
H2: Surface greenness and darkness will be positively
correlated.
H3: Surface permeability will be lower in areas with
green biolms.
H4: Sub-surface moisture levels will be higher beneath
green algal patches.
Measured greenness (a*) and hardness for
green and non-green areas
Measured greenness (a*) and darkness (L*)
on stone surfaces with patchy algal cover
Measured surface permeability for green (G)
and non-green (NG) areas.
Inferred sub-surface moisture levels for walls
with G and NG areas, before and after
experimental wetting

Objective 2: greening and environmental factors

Environmental factors: Green biolms are moisture-limited and H5: Green biolms will be most prevalent in cool, moist Measured greenness (a*) and possible
likely to be most dense in areas that trap and retain moisture locations on the surface of the stone (probably close to explanatory variables, including surface
e.g. rough, uneven and shaded surfaces or in regions with a ground level) and areas with a roughened texture. moisture, temperature and texture.
reliable supply of moisture e.g. close to ground level
154 N.A. Cutler et al. / Science of the Total Environment 442 (2013) 152164

year (Met Ofce, 2011), creating favourable conditions for the growth
of green algae. Models of future climate change predict increased sea-
sonality in the rst part of the twenty rst century, with wetter win-
ters and drier summers (Smith et al., 2010). The city is therefore the
ideal location to study the interaction of algal communities, the
stone surfaces on which they live and the environment.
We surveyed four sandstone heritage structures in central Belfast,
namely All Souls Church (AS), Crescent Church (CC), Ewart's Building
(EB) and Fitzroy Church (FC) (Fig. 1). All of the buildings were located
close to busy roads. The structures were similar in age (115
142 years old) and constructed from a variety of Carboniferous and
Permo-Triassic sandstones. The sandstones varied in terms of grain
size, texture and colour. Scrabo and Dugannon sandstones are light,
and range in colour from off-white to buff or pinkish in colour. They
are ne-grained and often contain laminations and lenses of silt and
clay, leading to a heterogeneous substrate on small spatial scales. In
contrast, Giffnock and Locharbriggs sandstones are biscuit-coloured
and red-brown, respectively. These sandstones are ne- to coarse-
grained and are usually well sorted (Curran et al., 2010). The wide
range of sandstone colours and textures present means that Belfast's
building stock may provide analogues for sandstone weathering in
many different locations.
2. Materials and methods
2.1. Sampling strategy
The greening of buildings is inherently patchy. Consequently, the
study was conducted at two scales, focusing on the properties of indi-
vidual blocks (centimetre scale) and larger sections of wall (metre
scale). In both cases, stone surfaces with obvious algal staining were
selected for study. Unpublished research by the authors suggests

Fig. 1. Details from the study structures, showing a) the ashlar walls of Ewart's building; b) detail of an ashlar block in Giffnock stone from the same building, showing G and NG
areas; the unevenness of the surface has been caused by the detachment of a ake of stone ~1.5 mm thick; c) the rock-faced walls of All Souls Church; note the inclined ashlar string
course; d) detail of a rock-faced, Scrabbo block from All Souls Church; note the uneven surface texture of the and patchy blackening. Green algal patches on are demarcated with
dotted lines and labelled G. The algal patches on All Souls Church are smaller and more dispersed than those on Ewart's Building.
 N.A. Cutler et al. / Science of the Total Environment 442 (2013) 152164 155

that the algal biolms on the study structures have similar species
compositions. For the block surveys, three to ve stone blocks were
purposefully selected on each of the four buildings (Table 2). Blocks
with patchy algal cover were selected so that green (G) and
non-green (NG) areas could be compared (Fig. 1b, d) and stone hard-
ness, permeability and surface colour were recorded. For the transect
surveys, vertical survey lines extending from the base of the wall to
~ 2 m above ground level were established on each building
(Table 3). The transect lines were located on north-facing sections
of wall with obvious algal staining. Surface moisture levels, surface
temperature and colour were recorded at 4 cm intervals along each
transect line (50 measurements/transect). Sub-surface resistivity
was measured via electrode arrangement at the same 4 cm intervals
along the transect lines. Stone hardness was also measured for each
block crossed by the transect lines.
2.2. Greening and biodeterioration
2.2.1. Surface hardness
Measurements of surface hardness were taken using an Equotip
type D portable hardness tester (Viles et al., 2011), which measures
hardness in Leeb units (Proceq SA, Zurich, Switzerland). For the
block surveys, ten measurements, concentrated in an area of ~ 1 cm 2
in the centre of the G and NG patches, were taken. The measurements
were then used to calculate mean hardness and the coefcient of var-
iation of surface hardness (CVSH), a normalised metric of variability
derived by dividing the sample standard deviation by the sample
mean. The assumption was made that the CVSH would act as a
proxy for the degree of weathering i.e. this metric would be higher
in weathered stone, due to the presence of softer, weathered patches
alongside less altered material. It is possible that a stone surface could
be highly weathered and uniformly soft (resulting in a low CVSH) but
none of the stone surfaces we surveyed exhibited this characteristic.
For the transect surveys, surface hardness was recorded in the centre
of each stone block on the transect line, again using 10 measurements
concentrated in an area of ~ 1 cm 2.
2.2.2. Appearance
The discolouration of stone surfaces was measured relative to an
unweathered surface of the same stone type. Colour measurements
based on the L*a*b* system were made using a hand-held Konica
Minolta CM-600d spectrophotometer (Konica Minolta Sensing Inc,
Osaka, Japan). Reference values were established on an unweathered
block of the same stone type, using the average of 10 measurements.
The study focussed on deviations on the red-green scale (a*) and on
the light-dark scale (L*), referred to hereafter as greening and
darkening. On these scales, negative values indicate that the
readings are greener and darker than the reference, respectively. In
L*a*b* colour space, the distance between two points (essentially a
vector length) is described as E. In principle, the untrained human
eye cannot detect a E value b 1 (although this does vary somewhat
according to hue, saturation and the sensitivity of the observer)
(Wyszecki and Stiles, 2000). Therefore, for an area of stone to be per-
ceptibly greener or darker than an unweathered reference sample, it
must have a a* or L*value b 1. For the stone blocks, deviations
from the reference values were calculated for the G and NG areas,
again using the average of 10 measurements. For the transect surveys,
single colour measurements were made at 4 cm intervals along the
transect line (i.e. 50 measurements/transect).
2.2.3. Movement of moisture
On the stone blocks, Karsten tube water penetration tests were
used to compare the permeability of G and NG areas. The Karsten
tubes comprised a 30 mm diameter hemispherical bulb with a
10 cm graduated tube attached. The open side of the bulb was ad-
hered to stone surface with a continuous compressible seal
(Blu-Tack) and 10 ml of water was added. The volume of remaining
in tube was recorded at 30 s intervals over a period of 15 min in
order to calculate inltration rate. Permeability tests were only
conducted on blocks with comparatively smooth, ashlar surfaces
that were suitable for the attachment of Karsten tubes. Inltration
rates for G and NG areas were compared with each other, and with
a reference rate established on a new wall of a similar stone type.
On the transect lines, sub-surface moisture conditions were in-
ferred for all ve buildings using 2D electrical resistivity tomography
(ERT). The ERT surveys were carried out with a GeoTom device
(Geolog 2000, Ausburg, Germany), following the methodology of
Sass and Viles (2006). Geoelectrical measurements were carried out
by applying a constant electrical current to the stone surface via elec-
trodes and measuring the resulting voltage difference at two poten-
tial electrodes (refer to Kneisel, 2003; Sass, 2003 for further details).
Resistivity values were then calculated from the known voltage and
current values. The device was connected via multicore cables to a
line of 50 self-adhesive ECG pads spaced at 4 cm intervals along the
transect lines (giving a total transect length of 1.96 m). We utilised
a Wenner electrode array, a conguration that has been used success-
fully in ERT on other buildings, as it provides good resolution of struc-
tures lying parallel to the stone surface and a high signal-to-noise
ratio (Sass and Viles, 2006, 2010). The resistivity measurements
were converted into geometrically accurate representations of
sub-surface resistivity using the Res2DINV inversion software with
robust inversion settings (Loke, 1999). The electrical resistivity of
building stone is largely determined by its moisture content (Loke,
1999); it was therefore assumed that low resistivity equated with

Table 2
Details of block survey (Sst = sandstone; RF = rock-faced, ASH = Ashlar).

Location Block no. Sst type Surface Aspect Property studied

Hardness Colour Permeability

All Souls Church (constructed 1896) 1 Scrabo RF W Y Y N

2 Scrabo RF W Y Y N
3 Scrabo RF W Y N N
4 Scrabo RF NW Y N N
5 Scrabo RF SW Y N Y
Crescent Church (constructed 1887) 1 Scrabo RF N Y N N
2 Scrabo RF N Y N N
3 Scrabo RF N Y N N
4 Locharbriggs ASH N Y Y Y
5 Locharbriggs ASH N Y Y N
Ewart's Building (constructed 1869) 1 Giffnock ASH N Y Y Y
2 Giffnock ASH N Y Y Y
3 Giffnock ASH N Y N N
Fitzroy Church (constructed 1874) 1 Locharbriggs ASH N Y Y Y
2 Locharbriggs ASH N Y Y Y
156 N.A. Cutler et al. / Science of the Total Environment 442 (2013) 152164
Table 3
Details of transect surveys.
Location Aspect Wall construction
All Souls Church N Coursed, rock-faced Scrabo Sst blocks with Doulting Lst string course
towards top of transect line;
Crescent Church N Squared and pitched, rock-faced Scrabo Sst blocks with dressed
Locharbriggs string course at top of plinth
Ewart's Building N Large, dressed Giffnock Sst blocks with projecting string course and
moulded plinth top
Fitzroy Church N Squared and pitched, rock-faced Scrabo Sst blocks with two dressed
Scrabo Sst string courses
high moisture availability and vice versa, although it is possible that
low resistivity values may also be due to the presence of salts, mate-
rial type and characteristics such as porosity.
After carrying out the initial ERT surveys, the movement of mois-
ture was assessed by means of a wetting experiment. Water was ap-
plied to the area in either side of the transect lines to simulate driving
rain, broadly following the methods of Sass and Viles (2010). First, 2 L
of water was applied in a ne mist spray over a period of approxi-
mately 5 min. The wall was wetted evenly in a swath extending ap-
proximately 10 cm either side of the transect line. ERT surveys were
then carried out 15 and 30 min after wetting to assess the ingress of
water into the wall. Resistivity-depth plots were produced by calcu-
lating the average resistivity values in 5 cm depth bins. The impact
of algal cover was assessed by comparing mean resistivity values in
zones below algal patches with those falling outside these zones.
The zone boundaries were dened by edges of the algal patches and
projected into the stone normal to the transect line.
2.3. Greening and environmental conditions
In addition to measuring surface hardness, colour and factors re-
lating to the movement of moisture, we also recorded environmental
parameters likely to be important in determining the distribution of
algal patches, including surface moisture, temperature and texture.
We restricted our analysis to the transect surveys. An analysis of co-
variance (ANCOVA) was used to model the relationship between
greening (a*) and the potential explanatory variables (moisture,
temperature, CVSH and surface texture).
2.3.1. Surface moisture and temperature
Surface moisture levels along the transect lines were measured
using a hand-held Protimeter moisture meter (GE, Faireld, CT,
USA). The device records surface moisture levels in % wood moisture
equivalent (w.m.e.); it is easy to use and routinely applied in assess-
ments of moisture in the construction industry and can be used on
stone to provide a picture of relative moisture trends (although, as
with ERT, readings may be inuenced by the presence of salts:
Eklund et al., 2012). Single Protimeter measurements were made at
4 cm intervals (in the same locations where the colour readings
were taken) and averaged for each block crossed by the transect
line (the number of measurements per block, n, varied according to
block size). Surface temperature measurements were made at the
same intervals, using a Fluke 62 mini infra-red thermometer (Fluke,
Everett, WA, USA). In the case of both moisture and temperature,
the focus in this study was on relative differences along the transect
line, rather than absolute values.
2.3.2. Surface texture
The blocks crossed by the transect lines varied in terms of surface
texture. For each sample point along the transect lines, surface tex-
ture was recorded as either ashlar (ASH, where the stone was worked
to a smooth, even surface) or rock-faced (RF, where the surface of the
stone was roughly hewn) (Fig. 1b, d).
Notes
Base of transect in shallow (c. 60 deep) trench; stone noticably
darkened with bright green and orange algal patches
Patchy blackening and green algal cover
Thick algal and bryophyte cover at base of plinth
Greening apparent across whole surface; stone crumbling in places;
building cleaned in 1980s
3. Results
3.1. Greening and biodeterioration
3.1.1. Physical integrity
In the block surveys, mean stone hardness varied widely from
308 91 to 546 62 Leeb units. There were no signicant differences
in hardness among the three stone types (ANOVA: F2,16 = 1.17, p =
0.33). CVSH ranged between 6 and 32% and was related to mean
hardness: harder surfaces were less variable. When data from all
the blocks were analysed, there was no signicant difference between
green (G) and non-green (NG) sections within blocks in terms of the
CVSH. However, the spread of the G data was heavily inuenced by an
outlying value (indicated by an arrow in Fig. 2). When this value was
omitted, G areas were signicantly less variable than NG patches
(t-test: t = 2.2, p = 0.045) as well as being signicantly harder
(t-test: t = 2.2, p = 0.04). The range in mean stone hardness was sim-
ilar in the transect surveys (315.1 27.5 to 391.5 16.8 Leebs). Stone
hardness exhibited considerable variability along the transect lines,
particularly on Fitzroy Church (Fig. 3d) where CVSH ranged from 7
to 58% (Table 4). There was no overall correlation between CVSH
and the degree of greening on the transect surveys (Table 5).
3.1.2. Appearance
Colour measurements made during the block survey revealed the
pronounced impact of algal staining: when G and NG areas were com-
pared, the mean difference in a* was 5.8. However, there was no
signicant difference in the darkening of G and NG areas on the
blocks (t-test: t = 0.71, p = 0.49). The results of the transect surveys
indicated that all of the surfaces were, on average, greener and darker
than unweathered stone of the same type (Table 4). Each of walls had
clearly dened green patches; All Souls Church also had patches of
positive a* values associated with a reddish-orange algal covering
Fig. 2. The relationship between mean hardness and CVSH for G (lled circles) and NG
(unlled circles) areas. The arrow indicates an outlying value from the G dataset.
 N.A. Cutler et al. / Science of the Total Environment 442 (2013) 152164 157

Fig. 3. A summary of the data collected from the transect surveys a) All Souls Church; note the positive peaks on the right hand side of the a* plot (which indicate patches of orange-red
algae), the relatively high moisture values close to ground level and the increase in temperature with increasing distance from the ground; b) Crescent Church; c) Ewart's building; note
the high variability in greening and moisture levels; d) Fitzroy Church; note the consistent greening and high variability in surface moisture and CVSH. The transect proles are shown on
the left hand margin of the plot (solid stone shaded grey). The vertical dotted lines on the a* and L* plots represent a departure of 1 units from the reference value derived from
unweathered stone; points to the left of these lines would be perceptibly greener or darker than unweathered stone. Error bars indicate 1 SE.

(Fig. 3a). The patches varied in size: those on the uneven, rock-faced large, continuous areas of green (Fig. 1a). The average level of greening
Scrabo walls (All Souls Church and Crescent Church) were small. In con- was less than the threshold value of 1 units on All Souls Church
trast, the smooth, ashlar walls of Ewart's Building were characterised by (probably due to the presence of red-orange algal cover), but more
158 N.A. Cutler et al. / Science of the Total Environment 442 (2013) 152164
Table 4
Summary of transect data (w.m.e. = wood moisture equivalent; SH = surface hardness; CVSH = coefcient of variation of surface hardness).
a* Mean moisture Moisture range
(% w.m.e.) (% w.m.e.)
All Souls Church 0.7 0.4 14 0.6 7.632.5
Crescent Church 1.5 0.2 15 0.5 7.229.8
Ewart's Building 4.4 0.7 20 1.4 8.564.5
Fitzroy Church 3.2 0.1 21 1.6 7.632.5
pronounced on the remaining buildings. Fitzroy Church was noticeably
green along the whole transect (Fig. 3d). The wall base was greener
than the upper parts of the transect lines on the Crescent Church and
Ewart's Building, but there was no discernible structure to the greening
on All Souls Church or Fitzroy Church. The darkening of the stone was
marked on all four structures, particularly All Souls Church (Fig. 3a).
The L* values recorded ranged from 2.3 to 29.6 but, as with
greening, there was little overall structure to the colour change:
the only gradient was apparent on Ewart's Building, which was
darker at the base (Fig. 3c). There was no signicant correlation be-
tween greening and darkening when all the blocks surveyed were
considered (Spearman-rank: Rs = 0.13, p = 0.42).
3.1.3. Movement of moisture
The results of the block analyses indicated that inltration rates
for weathered stone were higher than those for unweathered stone
for three of the four buildings (All Souls Church, Crescent Church
and Ewart's Building: detailed results not shown). Inltration rates
on the Fitzroy Church blocks were higher than unweathered stone
in NG areas and similar in G areas. There were no perceptible differ-
ences in inltration rate for G and NG areas for All Souls Church and
Crescent Church. For Ewart's Building and Fitzroy Church, inltration
rates were lower in G areas.
The ERT surveys conducted along the transect lines had a maxi-
mum penetration depth of ~ 32 cm. With the exception of Ewart's
Building, resistivity was broadly distributed according to a) height
above ground level and b) depth below stone surface. In terms of
height, the lower parts of the walls were dominated by low resistivity
values (cool colours i.e. blues and greens on the ERT plots: Fig. 4). In
terms of depth, three distinct zones were apparent: 1) a thin (~ 5 cm)
zone near the wall surface characterised by high resistivity values
(hot colours i.e. yellows and reds: Fig. 4); 2) a zone at intermediate
depth (~ 510 cm) characterised by low resistivity values and 3) the
deepest part of the wall, characterised by high resistivity values.
This pattern may be seen in the resistivity/depth plots (Fig. 5). For
All Souls Church, Crescent Church and Fitzroy Church (Fig. 5a, b and
d), mean resistivity increased with depth and the lowest resistivity
values were found at a depth of 510 cm. The opposite pattern was
observed for Ewart's Building: surface resistivity was variable but
generally high and mean resistivity decreased with increasing
depth. Across all the walls, the highest values (assumed driest)
tended to be clustered in high, deep locations; the lowest values
were generally in low locations of intermediate depth.
In addition to these broad patterns, it was also possible to discern
small-scale features, such as individual stone blocks and abrupt
changes in material (designated by Roman numeral in Fig. 4). For
Table 5
Correlations between the variables recorded on the transect surveys. Signicance
codes: * 0.05; ** 0.01; *** 0.001.
a* Surface moisture Surface temperature
Surface moisture 0.23
Surface temperature 0.37* 0.57***
L* 0.13 0.43** 0.28
CVSH 0.12 0.08 0.02
Mean temp. Temp. range L* Mean SH Mean CVSH
(C) (C) (Leeb units) (%)
11.5 0.2 9.613.4 19.5 0.6 391.5 16.8 24 2
12.5 0.1 10.613.6 12.3 0.5 366.4 24.6 19 3
9.5 0.04 8.810.0 8.4 1.4 361.1 16.8 20 3
10.5 0.1 9.411.4 11.5 1.6 315.1 27.5 27 4
example, stone blocks and mortar joints are clearly visible in the All
Souls plot (Fig. 4a(i)). Similar structures were much harder to discern
on the Ewart's Building plot, where the stone blocks were larger and
the intervening joints were thinner (Fig. 4c). Certain surface features
were associated with abrupt changes in resistivity. For example, the
lower string course (a stonework feature taking the form of a hori-
zontal band) on Ewart's Building had comparatively low resistivity
values (Fig. 4c(ii)). Inclined string courses with low resistivity were
also observed on the Crescent Church plot (Fig. 4b(ii)). However,
the upper string course on the Fitzroy Church (Fig. 4d(iii)) and the
string course on the All Souls plot (constructed of coarse, porous
Doulting limestone) had high resistivity (Fig. 4a(iii)).
Visual inspection suggested that green patches on the surface
were associated with relatively high resistivity values (i.e. inferred
drier conditions) in the intermediate subsurface. This was particularly
evident on Ewart's Building transect, where resistivity values in the
plinth (which had a thick algal coating) were higher than those in
the non-green block above (Fig. 4c(iv)). Similar patterns were ob-
served on the Crescent Church (Fig. 4b(iv)). Mean resistivity values
for the stone beneath G and NG zones provided some evidence to
support this observation. For example, mean resistivity values
below green patches were higher than those of adjacent non-green
areas for both All Souls Church and Ewart's Building (Fig. 6). The op-
posite was observed for the Crescent Church; however, high mean
values for NG areas were driven by exceptionally high resistivity in
a patch towards the top of the transect line. The association between
algal cover and high resistivity values was largely restricted to wall
bases. In the upper parts of the walls, where algal biolms were gen-
erally scarcer, there appeared to be no consistent relationship be-
tween greening and subsurface moisture levels.
The overall distributions of resistivity values did not change dur-
ing the wetting experiments. For example, wetting had almost no ef-
fect on the resistivity prole of the Ewart's Building transect, largely
because the applied water drained readily from the smooth surface
of the stone (the hydrophobic properties of extensive algal cover
may have promoted this process). Not surprisingly, the application
of water tended to lower resistivity values (Fig. 5a, cd). The impact
of wetting was apparent at all depths 15 min after the application
of water. The observed patterns were typied by the Fitzroy Church
section (Fig. 7). Prior to wetting (t = 0), the section was characterised
by high resistivity regions on the surface (Fig. 7(i)) and at depth,
mostly in the upper parts of the wall (Fig. 7(ii)). Individual blocks
were visible, including some with higher resistivity than their sur-
roundings (Fig. 7(iii)) and some with lower resistivity values
(Fig. 7(iv)). Fifteen minutes after wetting (t = 15), resistivity readings
at depth were generally lower (Figs. 5d, 7(v)). Surface resistivity was
also reduced in areas of high initial resistivity (Fig. 7(vi)). Drying at
the surface (Fig. 7(vii)) and at depth (Fig. 7(viii)) was evident
30 min after the application of water; toward the base of the wall, re-
sistivity was lower (Fig. 7(ix)). Overall, resistivity readings did not re-
turn to their original level in this period. Resistivity increased at depth
L* in some locations (for example, the deepest layers of the Ewart's
Building transect: Fig. 4c); anomalies of this type have been observed
in previous studies and have been interpreted as an artefact of the in-
version routine, rather than drying of the stone (refer to Sass and
0.15
Viles, 2010, for further details).
 N.A. Cutler et al. / Science of the Total Environment 442 (2013) 152164 159

Fig. 4. ERT surveys of four structures, showing the inferred 2D ERT section (LHS of each plot), a bar representing algal patches (shaded grey) and a plot of a* values at each electrode
position (electrodes spaced at 4 cm intervals along the transect). On the ERT sections, hot colours indicate high resistivity values (assumed dry conditions) and cool colours indicate
low resistivity values (assumed damp conditions). The surface of the wall is to the right of the section; the section starts at a depth of ~2 cm below the wall surface. Green patches are
demarcated where a* values are b1. The gures on the vertical axis of the ERT section indicate measured distances (in metres) along the transect. Key to areas indicated by roman
numerals: (i) individual stone blocks; (ii) string courses with comparatively low resistivity values; (iii) string courses with comparatively high resistivity; (iv) sub-surface areas with
anomalously high resistivity, relative to surroundings.

3.2. Greening and environmental conditions
3.2.1. Surface moisture and temperature
Mean surface moisture levels varied within a narrow range (14
0.6% to 21 1.6% w.m.e.), with considerable small- (centimetre-)
scale variability (Table 4; Fig. 3). Variation was greatest on Ewart's
building and Fitzroy Church (Fig. 3c and d). There was some evidence
that the wall bases were damper than higher sections (e.g. on All
Souls Church and Crescent Church: Fig. 3a and b). Surface tempera-
ture broadly followed this trend, increasing with height above ground
level in a linear fashion. On average, surface temperature varied by
2 C over the length of the transect lines (a vertical distance of
~ 2 m). On Ewart's Building, the temperature range was only 0.7 C;
variation was higher on the rock-faced walls, reaching 3.2 C on the
All Souls transect (Fig. 3a and d).
3.2.2. Correlation structure
There were some signicant correlations between the variables
recorded on the transect surveys (Table 5). There was a signicant
positive correlation between greening and surface temperature, indi-
cating that cooler surfaces were greener (although the relationship
was not signicant when the points with the two lowest temperature
values were removed: Fig. 8a). In addition, there was a highly signif-
icant negative correlation between surface moisture and surface tem-
perature (moist surfaces were cooler); moisture levels were also
related to darkening (drier surfaces were darker) (Fig. 8b and c). A
simple linear regression of greening on temperature was signicant
(p = 0.02) and indicated that temperature accounted for 14% of the
variance in a*. None of the other potential explanatory variables
was signicant in the ANCOVA. There was no overall correlation
between greening and surface moisture; these variables followed
each other closely on Ewart's Building (Fig. 3c), but the correlation
was not signicant. Greening and darkening were correlated on
Ewart's Building and the Crescent Church (green areas were darker),
but not on the other walls. There was no signicant difference between
the degree of greening on rock-faced and ashlar blocks (MannWhitney
U-test, p = 0.36). None of the correlations with CVSH were signicant.
4. Discussion
The surveys demonstrated that exposed stone surfaces that had
been exposed in Belfast for around 100 years were, on average,
greener and darker than recently-exposed stone of the same type.
The link between greening and the physical condition of the stone
was less clear and it is likely that the physical disintegration of
stone in Belfast is mainly driven by abiotic factors. However, there
was evidence that hardness was less variable in green areas. Thin
biolms on the upper parts of walls (sampled in the block survey)
had little impact on permeability. However, the ERT surveys
suggested that thick algal biolms on the bases of the walls were as-
sociated with relatively dry subsurface conditions. Greening was
poorly correlated with the environmental variables measured in this
study, although there was a signicant negative correlation between
temperature and greening, suggesting a link between evaporation
rates and biological activity.
4.1. Physical structure [H1]
Harder surfaces exhibited less variability in hardness than softer
surfaces (Fig. 2). There was no compelling evidence to link green
160 N.A. Cutler et al. / Science of the Total Environment 442 (2013) 152164
Fig. 5. Resistivity depth plots for a) All Souls Church; b) Ewart's Building and c) Fitzroy Church. The mean gures are derived from resistivity values binned at 5 cm depth intervals.
The values plot for t = 0 indicates the values under ambient conditions; the t = 15 and t= 30 plots indicate the impact of wetting. Note the differences in the scaling of the y-axis.
Error bars indicate 1 SE.
biolms to high levels of weathering. Conversely, there was evidence
from the block studies that algal patches were associated with less
weathered surfaces (i.e. harder patches with lower CVSH). This
might indicate that green algal cover has a broadly bioprotective
role. Of course, green algal patches can be transient and the biolms
surveyed in this study may not have had sufcient time to contribute
to weathering processes. Detailed examination by scanning electron
microscope (SEM) would be required to denitively establish wheth-
er or not microbes were associated with weathering (De Los Ros et
al., 2004). However, it seems likely that the main factors driving the
weathering of stone in the study locations were abiotic processes.
Fig. 6. Comparison of mean resistivity values in stone lying beneath green algal patches
(G) and areas without algal cover (NG). Error bars indicate 1 SE.
Buildings in Belfast city centre are exposed to a range of abiotic dete-
riorative processes, notably salt weathering (Turkington and Smith,
2000). Although concentrations of atmospheric SO2 and NOx are in
decline (DOE, 2005), air quality in the city can be poor due to a com-
bination trafc congestion, meteorological and topographic factors
(Smith et al., 2010). The only way to denitively establish the relative
contribution of greening to the physical deterioration of stone would
be the use of long-term, high resolution studies on suitably cong-
ured test walls.
4.2. Appearance: greening and darkening [H2]
The study identied changes in the appearance of the stone sur-
faces over time. On average, the stone surfaces studied were greener
and darker than fresh stone of the same type. Darkening of stone can
occur for several reasons, including the gradual accumulation of dust
particles and/or the formation of pigmented microbial biolms (in-
cluding crusts of lichenized fungi e.g. Verrucaria spp.) Microbial action
might also induce the oxidation of manganese in the stone (de la
Torre and Gomez-Alarcon, 1994), which would also lead to darken-
ing, although we did not investigate this. Greening and darkening
appeared to be closely associated in plinth areas. However, the
lower parts of a wall are prone to soiling (e.g. by particles of soil
entrained in rain splash) so the darkening of the stone in these loca-
tions is not necessarily related to biological processes. The relation-
ship between greening and darkening was weaker on the upper
parts of the wall: in the block surveys, for example, there was no cor-
relation between algal cover and darkening, despite stark differences
in greenness between G and NG areas. We did not distinguish be-
tween biotic and abiotic darkening. However, it seems likely that
the darkening of the stone was driven by processes that have
 N.A. Cutler et al. / Science of the Total Environment 442 (2013) 152164 161

Fig. 7. The results of the wetting experiment carried out on the Fitzroy Church transect. Time after wetting is indicated in minutes (i.e. t= 0, 15 and 30 min). Key to areas indicated
by roman numerals: (i) high surface resistivity; (ii) high resistivity at depth; (iii) individual block with high resistivity; (iv) individual blocks with lower resistivity values than their
surroundings; (v) lowered resistivity at depth (at t = 15); (vi) lower surface resistivity (at t = 15); (vii) evidence of surface drying (increased resistivity, at t = 30); (viii) increases
in resistivity at depth towards top of wall (at t= 30); (ix) decreased resistivity at depth towards base of wall (at t = 30). Note the resistivity scale differs from Fig. 4. Error bars
indicate 1 SE.

operated independently from biolm formation. In Belfast, as in many
other urban areas, blackened stone is often associated with the for-
mation of surface crusts that form when minerals in the stone react
with atmospheric pollutants such as SO2 (Turkington and Smith,
2000). Anecdotal evidence suggests that green biolms are associated
with black crusts (Smith et al., 2010), possibly because changes in
stone properties associated with crust formation increase the
bioreceptivity of the substrate. This was observed on All Souls Church
(Fig. 1 d). However, we could not nd a consistent relationship when all
of the buildings were considered. Whilst greening and darkening may
coincide, it seems likely that this is due to a confounding variable
(such as moisture supply) rather than a direct causal correlation.
4.3. Moisture movement [H3]
The block studies indicated that the permeability of the old stone
was higher than that of new stone, suggesting an opening-up of the
stone structure by weathering processes. However, when a larger
number of individual blocks were studied, permeability was not con-
sistently related to algal cover. The lack of signicant variation may be
related to the thinness of the biolms studied; in each case the algal
covering on the blocks was bright green but thin enough so that the
colour and texture of the underlying stone could be seen. Much
thicker algal coverings were observed on some of the transect studies
(e.g. at the base of the Ewart's Building transect) and these seem to
162 N.A. Cutler et al. / Science of the Total Environment 442 (2013) 152164
Fig. 8. Scatterplots illustrating signicant correlations in the dataset, based on block averages from the transect lines: a) relative greening (a*) and temperature; b) surface mois-
ture and surface temperature and c) relative darkening (L*) and surface moisture.
have had more related to the properties of the stone (surface temper-
ature and moisture, in particular). It may be that there were insuf-
cient algal cells present on the surface of the individual stone blocks
studied to affect permeability or initiate signicant bioweathering.
The ERT analyses revealed details of the 2D distribution of mois-
ture in the survey walls. On average, the lowest mean resistivity
values were recorded for Fitzroy Church; this was also the greenest
building, suggesting a connection between moisture and greening.
There was also evidence from the ERT surveys that the lower parts
of the walls studied were both damper and greener than the upper
sections. This result is not surprising, as the lower parts of walls are
subject to the capillary rise of ground water and rain-splash.
There was evidence that algal cover was associated with higher re-
sistivity values (and inferred drier conditions) at depth (see, for exam-
ple, Fig. 6). This pattern was apparent in the ERT data from All Souls
Church and Ewart's Building. It was also apparent on the ERT plot for
Crescent Church, although a patch of very high resistivity values
meant it was not observed in averaged values. It is therefore possible
that algal biolms impede the ingress of moisture, consistent with ear-
lier studies (Smith et al., 2011). However, it will require systematic,
long-term monitoring of subsurface moisture levels to determine
whether or not algal biolms are responsible for this effect and what
(if any) impact this has on the long-term stability of stone structures.
In the wetting experiments, mean resistivity values did not return
to their original values within 30 min and it was difcult to assess the
impact of algal cover on moisture levels at depth. However, there was
evidence of rapid surface drying and it is likely that, outside of periods
of prolonged rainfall, the greater part of the wall surface is dry for
much of the time. Under these conditions, metabolic activity in algal
cells is likely to be restricted to comparatively short spells following
wetting events. If this is the case, subsequent studies will have to
use continuous measurements of surface moisture and integrate the
results over time, to derive a time-of-wetness metric. These future
studies would also have to establish the minimum moisture threshold
above which lithobiotic algae can function.
4.4. Environmental parameters
If moisture was the main determinant of greening, then the patchy
distribution of the algae, coupled with the sharp patch boundaries,
would suggest signicant and abrupt variability in surface and
near-surface moisture levels. This was not observed: surface moisture
values were highly variable, but largely unrelated to the incidence of
algal patches. This result was surprising, as other studies have indicat-
ed that moisture is a key limiting factor for sub-aerial algae
(Bellinzoni et al., 2003).
The weak correlation between surface moisture levels and the in-
cidence of algal patches may have been due to the survey strategy
adopted. Surface moisture levels on exposed stone surfaces are ex-
tremely dynamic, varying on a variety of timescales from minutes to
months (Sass and Viles, 2006). The measurements taken in this
study were point values and cannot reveal long-term patterns. How-
ever, moisture measurements performed during comparatively dry
spells can reveal areas in which residual moisture persists, and it is
these areas that are likely to be most favourable for algal growth. Sim-
ilarly, point measurements of temperature can reveal trends in tem-
perature that could drive differences in evaporation rates, and
hence available moisture. It is possible that the moisture dynamics
that inuence algal distribution cannot be captured by point studies.
Furthermore, it may be that the Protimeter, whilst suitable for rapid
surveys, is a poor guide to the moisture conditions experienced by
the algae. The presence of dissolved salts in pore water, for example,
can produce anomalously low resistivity readings in geoelectrical de-
vices (Sass, 2003). Even though the walls we studied showed no ob-
vious signs of eforescence, it could be that the presence of
sub-surface salts confounded our results.
In contrast to moisture readings, surface temperature was correlated
with greening. Areas with lower temperatures were, on average,
greener than warmer areas. The relationship was strongest on walls
where the plinth was very green; temperature performed less well as
a predictor of greening on the upper parts of walls, where temperatures
 N.A. Cutler et al. / Science of the Total Environment 442 (2013) 152164
were generally higher. Abrupt changes in temperature associated
with discrete algal patches were not observed. The overall range of
temperature variation was comparatively small (~ 2 C) and proba-
bly insufcient to make a signicant impact on algal metabolism.
The relationship between greening and temperature was therefore
probably due to lower rates of evaporation in cooler areas at the
base of walls, although other effects in these areas, e.g. an increased
supply of moisture and nutrients through rain-splash, may also have
been important.
4.5. Unmeasured environmental variables
The greening observed on the transect lines exhibited high spatial
heterogeneity, often on sub-block scales, but was unrelated to most of
the environmental variables recorded. The weak relationship be-
tween greening and the explanatory variables meant that it was not
possible to satisfactorily model the greening of sandstone buildings
in Belfast. Of the environmental variables studied, only surface tem-
perature was related to greening, and this factor only accounted for
around 14% of the observed variability in greening. This indicates
that there were other, unmeasured, factors driving the incidence of
algal patches. It is likely that a recongured metric of moisture, e.g. an-
nual time-of-wetness, would improve the explanatory power of the
model. Other likely candidates for explanatory variables include ne-
(sub-block-) scale surface morphology and macronutrient status.
Surface morphology encompasses centimetre- and sub-centimetre
surface roughness and inclination (measured relative to a vertical
plane). On the rock-faced Scrabo walls, algal patches appeared to be as-
sociated with centimetre-scale variations in surface topography. Certain
microtopographic congurations e.g. hollows or shallow slopes, could
be more suited to algal colonisation because they collect moisture
and/or dry more slowly (Miller et al., 2012). At a larger scale, the incli-
nation of architectural features may inuence hydrological regimes. Ob-
servations of buildings in Belfast suggest that inclined stone surfaces
(e.g. the tops of window sills, buttresses and projecting string courses)
are preferentially colonised by algae, perhaps because they dry more
slowly than vertical faces. The underside of projecting features may at-
tract algae because they are often in shade. The simple categorisation of
surface texture used in this study did not capture differences in green-
ing related to surface morphology. However, it is possible that high res-
olution studies that focus on ne-scale surface morphology (perhaps
systematically contrasting inclined and vertical features) will be more
successful. Exposure trials utilising stone blocks with a variety of archi-
tectural features would be particularly useful.
Macronutrient supply is a key factor regulating biological activity
in all ecosystems and spatial heterogeneity in this factor is often asso-
ciated with patchiness at various scales. Building stone is an oligotro-
phic habitat and nutrients such and nitrogen (N) are likely to be
limiting or co-limiting (with moisture) to algal productivity (John,
1988; Round, 1981). Nitrogen is unlikely to be derived directly from
stone substrates; N will therefore be supplied predominantly through
wet and dry deposition. Changes in atmospheric concentrations of
NOx are therefore likely to inuence overall levels of greening and
may be more inuential than changing precipitation patterns in this
context. However, small-scale hydrological factors are also likely to
inuence the distribution of deposited macronutrients across the
face of buildings, and hence the distribution of algal patches. This
phenomenon has not been studied, to our knowledge, but it could
certainly be a contributory factor to ne-scale distribution of algal
greening.
In addition to environmental conditions, biological processes
e.g. dispersal or competition for limiting resources, can drive spatial pat-
terning in biological communities (Fortin and Dale, 2005). Species-level
differences in the dispersal capability, or the distribution of parent
populations, could lead to local differences in the composition of algal
communities. It is difcult to generalise about this factor without
163
knowing the detailed species composition of the biolms studied. The
airborne dispersal of green algae is poorly understood (Sharma et al.,
2006, 2007) and the complex wind environments of urban settings
make it difcult to estimate how dispersal might impact on the distribu-
tion of algal communities. However, the microbes found on building
stone tend to be generalists and green algae are found in a host of
terricolous, corticolous, saxicolous and terricolous habitats, including
other buildings (Rindi, 2007; Rindi and Guiry, 2003, 2004). Source
areas for colonisation are therefore likely to be numerous. Given the
geographical proximity of the study structures, and similarities in their
surroundings, there was nothing to suggest that dispersal differences
contributed to the observed variations in greening.
In addition to dispersal, other biological processes might also have
structured the biolms in this study. Community interactions are poorly
understood for microbes in the environment and exceptionally difcult
to monitor. It is not clear whether macroscale patterns (such as large
algal patches) can emerge from microscale competitive interactions.
Contagious spread following a random colonisation event could lead
to patchiness that would not necessarily be related to environmental
variables, providing these factors were not limiting. Fortuitous colonisa-
tion events might be favoured by surface irregularities, as discussed
above. Finally, there is likely to be a substantial random component to
the incidence of algal biolms, e.g. cleaning or abrasion of the stone sur-
face, overshadowing by trees or buildings and transient moisture
sources (e.g. leaks), to name a few.
Variations in community composition could impact on studies of this
type, as different communities (with different rates and types of meta-
bolic activity) would reduce the comparability of different structures.
Unpublished research by the authors suggests that the algal communi-
ties on Belfast sandstone are very similar in composition. However,
other studies of this type would have to establish algal community com-
position (including an assessment of whether the community included a
signicant component of cyanobacteria) before comparing rates and
patterns of weathering between buildings.
5. Conclusions
Green biolms can have an aesthetic impact on heritage struc-
tures, but in our study this appeared to be limited to green staining
(i.e. the algae were not consistently associated with the darkening
of the stone). Whilst the growth of green algal biolms may be con-
sidered visually detrimental, we could nd no evidence that algal
greening is directly involved in the weathering of sandstone (other
techniques, e.g. use of scanning electron microscopy, would be re-
quired to establish denitively whether or not green algae contribut-
ed to the weathering of the stone). ERT data suggest that algal patches
are associated with higher resistivity values (and, by implication,
lower moisture levels) in the immediate subsurface. Coupled with
data on the relationship between stone hardness and algal cover,
there is therefore evidence that green algae might have a broadly
bioprotective role. Further study, ideally based on test structures,
will be required to establish whether or not this is the case. Our at-
tempts to model greening suggest that surface temperature is corre-
lated with the distribution of algae across the surface of a stone
building, but this factor is likely to be confounded with other vari-
ables. It is likely that moisture is a key factor in determining the over-
all distribution of green algal biolms. However, the relationship
between moisture levels and greening is not straightforward and is
likely to be heavily dependent upon the temporal distribution of
moisture and, possibly, transient wetting events. Overall, it seems
that algal greening of sandstones is more related to environmental
conditions (particularly climate and atmospheric chemistry) than
stone type. It may therefore be possible to model algal greening of
sandstones from the scaled-down outputs of regional climate
models.
164 N.A. Cutler et al. / Science of the Total Environment 442 (2013) 152164
Acknowledgements
We thank three anonymous reviewers for their insightful com-
ments. This work was funded by EPSRC grant no. EP/G011338/1
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