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 Israel Journal of Ecology & Evolution

ISSN: 1565-9801 (Print) 2224-4662 (Online) Journal homepage: http://www.tandfonline.com/loi/tiee20

Green roofs: what can we learn from desert

plants?

Inga Dirks, Buzi Raviv, Oren Shelef, Amber Hill, Amir Eppel, Moses Kwame
Aidoo, Brian Hoefgen, Tal Rapaport, Hila Gil, Endale Geta, Amnon Kochavi,
Itay Cohen & Shimon Rachmilevitch

To cite this article: Inga Dirks, Buzi Raviv, Oren Shelef, Amber Hill, Amir Eppel, Moses Kwame
Aidoo, Brian Hoefgen, Tal Rapaport, Hila Gil, Endale Geta, Amnon Kochavi, Itay Cohen &
Shimon Rachmilevitch (2016) Green roofs: what can we learn from desert plants?, Israel Journal
of Ecology & Evolution, 62:1-2, 58-67, DOI: 10.1080/15659801.2016.1140619

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Israel Journal of Ecology & Evolution, 2016
Vol. 62, Nos. 1 2, 58 67, http://dx.doi.org/10.1080/15659801.2016.1140619

Green roofs: what can we learn from desert plants?

a
Inga Dirks , Buzi Raviv , Oren Shelefa, Amber Hilla, Amir Eppel b, Moses Kwame Aidoo a, Brian Hoefgena, Tal
a

Rapaporta, Hila Gila, Endale Getaa, Amnon Kochavi a, Itay Cohena and Shimon Rachmilevitcha*
a
French Associates Institute for Agriculture and Biotechnology of Drylands, Jacob Blaustein Institutes for Desert Research, Ben-Gurion
University of the Negev, Beersheba, Israel; bInstitute of Plant Sciences, Agricultural Research Organization, Gilat Research Center,
Beersheba, Israel
(Received 9 October 2014; accepted 17 December 2015)

Green roofs in the Mediterranean region are often exposed to high levels of radiation, extreme temperatures, and an
inconsistent water supply. To withstand these harsh conditions in shallow soils and poorly aerated growth media, plants
must be armored with adaptations. Strategies that have evolved in desert plants can play significant roles in the use of
plants for green covers. In the following, we will specifically focus on (1) heat and radiation, (2) drought, and (3) salinity.
Further, we will discuss (4) interactions between neighboring plants. Finally, we will (5) propose a design for diverse green
roofs that includes horticultural and medicinal products and provides diverse habitats. Many desert plants have developed
morphological and anatomical features to avoid photo-inhibition, which can be advantageous for growth on green roofs.
Plants exhibiting C4 photosynthesis or crassulacean acid metabolism (CAM) photosynthesis have a protected hydraulic
system that enables growth under dry conditions. Furthermore, dew and high levels of relative humidity can provide
reliable water sources under limited precipitation. Halophytes are protected against salinity, ionic specific stress, and
nutritional imbalances, characteristics that can be advantageous for green roofs. Under limited space, competition for
resources becomes increasingly relevant. Allelopathy can also induce the germination and growth inhibition of
neighboring plants. Many desert plants, as a result of their exposure to environmental stress, have developed unique
survival adaptations based on secondary metabolites that can be used as pharmaceuticals. A systematic survey of plant
strategies to withstand these extreme conditions provides a basis for increasing the number of green roof candidates.
Keywords: extensive green roofs; urban ecology; urban biodiversity; green roof design; desert plants; built environments

Introduction green roofs (Brenneisen 2003; Brenneisen 2004; Darius &

Green roofs are composed of vegetation, a growth
medium and structures that provide various environmental
benefits for urban areas, which often lack plant cover and
recreational space. These benefits improve the buildings
energy consumption by regulating biogeochemical cycles
such as thermoregulation, hydrological processes, and gas
exchange (Del Barrio 1998; Oberndorfer et al. 2007).
Significantly, lower air and surface temperatures were
measured when the roofs and walls of street canyons were
covered with green biomass across different climatic
zones (Alexandri & Jones 2008). For example, lawns on
green roofs decreased indoor temperatures in the range of
2 4  C throughout the day, thus mitigating the need for
air conditioning with its associated expense (Liang &
Huang 2011). By the assimilation and fixation of gaseous
pollutants, such as ozone (O3), nitrogen dioxide (NO2),
and sulfur dioxide (SO2), plants growing on green roofs
can act as air purifying agents (Currie & Bass 2008). In
addition, green roofs can restrain surface runoff in urban
areas, thus reducing the input of water loads into sewage
systems (K ohler et al. 2001; Mentens et al. 2006). Further-
more, green roofs can substantially impact local biodiver-
sity by providing undisturbed habitats. For instance, both
high levels of diversity and a proportion of endangered
plant and insect species have been found on extensive
Drepper 1984).
Plant growth on green roofs occurs under artificial
conditions that differ from natural ecosystems in several
ways. As a first step, we present the physiological adap-
tions of plants native to the Negev Desert that cope with
extreme (1) heat and radiation, (2) drought, and (3) salin-
ity. In addition, knowledge about plant interactions has
been generated that might hold substantial information for
the creation of diverse and persistent communities. Thus,
as a second step, we review (4) interactions between
neighboring plants. Based on the physiology of desert
plants, we propose (5) a roof design for rooftop gardens
that provides both perennial cover and diverse habitats.
The benefits that can be obtained from the different spe-
cies presented as potential green roof candidates are sum-
marized in Table 1.
(1) High radiation and extreme temperatures.
Extreme levels of heat and radiation dominate the
green roof environment, especially during sum-
mer months. Average maximum air temperatures
during the summer months in Tel Aviv reach
between 27.8 and 31.1  C (Cohen 2013). Whereas
air temperatures exceed 30  C during the summer
months, ambient temperatures on roofs and walls

*Corresponding author. Email: rshimon@bgu.ac.il

2016 Informa UK Limited, trading as Taylor & Francis Group

Table 1. Benefits and survival strategies of potential green roof candidates.

Ecosystem service Life form Survival strategy Species Origin Reference

Perennial roof cover Perennial, shrub Bright waxy leaves avoid Atriplex halimus Middle Eastern desert Streb et al. (1997)
photo-inhibition
Perennial roof cover Perennial shrub Hairy leaves avoid photo- Encelia farinos North American deserts Ehleringer et al. (1976)
inhibition
Medicinal properties Perennial shrub Dew uptake Artemisia sieberi Irano-Turanian region Hill et al. (2015), Friedjung et al. (2013),
allelopathic effects Mohamed et al. (2010), Ghazanfar
(2012)
Medicinal properties, Perennial shrub Dew uptake Artemisia judaica Southwestern regions of Friedjung et al. (2013), Putievsky et al.
allelopathic effects the Middle East, (1992)
eastern Sahara
Medicinal properties, Perennial shrub Origanum dayi Northern Negev and Friedjung et al. (2013), Danin (1983),
allelopathic effects Judean Deserts, Israel Dudai et al. (2003)
Fragrant flowers are attractive to Perennial shrub C4 perennial shrub Tidestromia oblongifolia, Central American deserts Bjorkman et al. (1972)
bees, butterflies, birds
Esthetics, nectar for honey bees Perennial shrub CAM photosynthesis, Lampranthus blandus South Africa Ben-Dov et al. (1993), Ben-Dov et al.
halophyte (2001), Choukr-Allah et al.(1996)
Esthetics, perennial roof cover Perennial shrub CAM photosynthesis, Drosanthemum hispidum South Africa Pittenger (2001)
halophyte
Antibacterial and antidiarrheal, Perennial grass C4 grass Cymbopogon citratus South Asia, Southeast Shah (2011), Tzortzakis and Economakis
antioxidant compounds, high Asia (2007)
productivity
Essential oil content Perennial grass C4 Cymbopogon parkeri Saharo-Arabian region Avoseh et al. (2015), Vogel et al. 1986
Israel Journal Of Ecology & Evolution

Roof cover on diverse roofs Summer annual C4 Eragrostis barrileri Mediterranean and Saharo Al Harthi et al. (2008), Vogel et al. 1986
during dry summer months grass Arabian regions
Roof cover on diverse roofs Summer annual C4 Setaria virides Euro-Siberian, Irano- Petti et al. (2013), Vogel et al. 1986
during dry summer months grass Turanian and
Mediterranean regions
Temperature insulation, Annual and Shift between C3 and CAM Sedum ssp Klett et al. (2012), Van Woert et al. (2005),
perennials have nursery effect perennial photosynthesis, high Werthmann (2007)
for annuals drought tolerance, shallow
root system
Esthetics Annual Low capacity of enzymatic Anastacia hierochunta Middle Eastern deserts Eppel et al. (2014)
photo protective processes
Esthetics Annual Dew uptake Salsola inermis Middle Eastern deserts Barazani and Golan-Goldhirsh (2009), Hill
et al (2015)
59
60 I. Dirks et al.
often reach much higher values. For example,
temperatures of 60  C were measured on southern
exposed walls in Shanghai in comparison to an
average ambient air temperature of only 28.6  C
during the summer months (Shen 2011). The tem-
peratures of the synthetic membranes used for
green roof construction can reach up to 80  C.
Stonecrop (Sedum ssp.) leaf temperatures reached
up to 60  C during the summer in 2009 on roof
surfaces in New York City (Gaffin 2010). The
average maximum air temperature during this
summer in New York was 25.6  C (National Oce-
anic and Atmospheric Administration, NOAA,
National Centers for Environmental Information).
(2) Water shortage. Plants on green roofs are exposed
to frequent desiccation because shallow soils dry
out very easily (Getter & Rowe 2006), and plants
are characterized by high levels of transpiration.
In C4 plants, CO2 can be fixed under very low
concentrations (Bowes 1991; OLeary 1982),
which allows the assimilation of CO2 with a small
stomatal aperture, thus leading to higher photo-
synthetic water use efficiency (Kocacinar & Sage
2003; Gowik & Westhoff 2011). Furthermore,
plants exhibiting crassulacean acid metabolism
(CAM) photosynthesis are extremely tolerant to
drought (Sayed 2001).
In addition, water sources other than precipitation might
be relevant. Dew and high relative humidity can be very
reliable water sources in contrast to the high variability of
precipitation events in semiarid areas. Plants with the ability
to use dew as a water source for growth and photosynthesis
could be an excellent choice for use in the cultivation of
extensive green roofs. In the Negev Desert, moisture uptake
by plants was positively correlated with the dew accumula-
tion on leaf surfaces in the two species Salsola inermis
Forssk and Artemisia sieberi Bess (Hill et al. 2015).
(1) Salinity. High radiation, which results in high
potential evapotranspiration, causes plants to face
salinity stress. Salinity entails osmotic stress,
ionic-specific stress, and nutritional imbalance
(Greenway & Munns 1980). Halophytes are plants
that are specialized in salinity tolerance. They rep-
resent only 2% 3% of all plant species, and they
succeed in conditions that are harmful to non-hal-
ophytes (Waisel 1972; Glenn et al. 1999).
(2) Interactions between neighboring plants. The area
of a typical green roof is limited by the boundaries
of the buildings roof area and the capacity of the
roof to carry soil, i.e., the maximum depth of the
soil on a given roof area. Plant growth in shallow
soils might be exposed to resource limitation and
to intensified interactions between plants.
(3) Diverse and sustainable garden design. For the
establishment of diverse and sustainable green
roofs, heterogeneous habitats with varying light
conditions and soil depths are fundamental. In
addition, the use of both annual and perennial
plants can help to increase diversity. Resource
competition and biotic interactions need to be
taken into consideration with respect to the fact
that they change with environmental conditions.
Based on physiological factors, we propose a number
of plants mostly native to the Negev Desert that have the
ability to grow on green roofs, therefore helping to main-
tain biodiversity in urban environments. The persistence
of wild populations in urban areas can contribute to biodi-
versity enrichment, in light of the fact that human-domi-
nated landscapes will not revert to wilderness anytime
soon (Ehrlich & Pringle 2008).
How can the radiation tolerance of desert plants
support green surfacing in urban areas?
Plant photosynthesis depends on solar radiation. From the
total radiation that reaches the earths surface, plants can
only use the radiation in the visible spectrum for their
photosynthesis (Zhu et al. 2008). Levels of solar radiation
in deserts tend to be high due to their geographical loca-
tions and the low cloud coverage in these areas. The aver-
age solar radiation in the Negev is 7 8 kWh/m2 day-1,
while in regions with less radiation, such as central
Europe, the radiation level is less than half of this value
(http://solargis.info/).
When plants are exposed to an excessive amount of
light, surpassing what they are able to use in photosynthe-
sis, a state of photo-inhibition may occur (Takahashi &
Murata 2008). Photo-inhibition is a light-dependent
reduction of plant photosynthetic capacity, leading to inhi-
bition of plant growth (Long & Humphries 1994). In order
to avoid photo-inhibition, plants have evolved anatomical,
biochemical, and enzymatic features and processes in
order to deal with the potential damage of high light inten-
sity (Murchie & Niyogi 2011). Such processes will be
fundamental for plant performance on green roofs.
In many desert plants, especially in perennials, adapta-
tion to a high level of radiation involves features and
mechanisms that reduce the amount of absorbed radiation,
thereby avoiding potential photo-inhibition and reducing
heat and water loss (Ehleringer & Mooney 1978;
Ehleringer 1981). Two examples of these features that
assist in avoiding photo-inhibition are the waxy bright
leaves that occur in the species Atriplex halimus L., com-
mon in Middle Eastern deserts (Streb et al. 1997), and the
leaf hairs (Pubescence) that are found in the species Ence-
lia farinosa Torr. & Gay, which grows in the deserts of
North America (Ehleringer et al. 1976). Admittedly, the
abovementioned adaptation methods can reduce the
plants capacity for photosynthesis, but they also confer
photo protection to the plant under high levels of radia-
tion, which is essential for physiological activity under
these extreme conditions. Removing waxes, for example,
resulted in a significant decrease in the ratio of variable to
maximum fluorescence (Fv/Fm ratio) and, in severe cases,
damage from photo-inhibition as indicated by an increase
in the initial fluorescence (F0) (Robinson et al. 1993).
High radiation levels in deserts lead to an increase in
 Israel Journal Of Ecology & Evolution
temperature and water evaporation in both plants and soil.
Therefore in many cases, desert plants adaptation to high
radiation overlaps, at least partially, with their adaptations
to water scarcity and high temperatures (Rewald et al.
2012). The coexistence of adaptations to drought and
intense radiation enables the survival and physiological
activity of perennial plants on green roofs.
In contrast to many perennial desert species, some
winter annual species deal with a high amount of radiation
by exhibiting high photosynthetic activity, for example
Anastatica hierochuntica L., the Rose of Jericho. A. hiero-
chuntica is an annual desert plant, found in the Middle
East, that has a high capacity for photosynthesis and high
transpiration rates, accompanied by a low capacity for
enzymatic photo protective processes (Eppel et al. 2014).
This is an example of a winter annual desert plant that
uses the short time period during the winter months when
sufficient soil water is available in order to grow quickly
and reproduce seeds before the arrival of summer.
Various responses of potential green roof candidates to
water stress
Plants native to arid and semiarid regions can be character-
ized by various strategies with which they cope with water
scarcity, while maintaining relatively high productivity.
In C4 plants, CO2 can be fixed under very low concen-
trations in the mesophyll cells due to the high abundance of
phosphoenolpyruvate carboxylase (PEPCase) and other
enzymes that concentrate CO2 around the enzyme ribulose-
1.5-bisphosphate carboxylase/oxygenase (OLeary 1982;
Bowes 1991; Sage 2004). The small stomatal aperture
decreases transpiration and allows photosynthesis to be sus-
tained under drying atmospheric and soil conditions (Obern-
dorfer et al. 2007). An analysis of the subtypes of C4 plants
revealed a higher presence of aspartate-forming NAD-malic
enzyme grasses under extremely xeric conditions, whereas
malate-forming NADP-malic enzyme grasses occurred
when water stress was less severe (Vogel et al. 1986).
Most C4 plants in Israel belong to the Gramineae
grasses and Chenopodiaceae shrubs (Frumkin et al. 2000),
which have recently been incorporated into the family of
Amaranthaceae. The Judean Desert and the Central Negev
Highlands have a relatively low proportion of C4 plants,
due to the fact that precipitation and low temperatures
synchronize during the winter months. In these regions, a
high correlation between decreasing precipitation and the
presence of C4 plants was found (Vogel et al. 1986). The
Arava Valley and the coastal plains along the Gulf of
Suez have warm temperatures during the winter months
and show a significantly higher proportion of C4 grasses,
which constitute up to 70% 100% of all grasses (Vogel
et al. 1986).
Information about the life cycle, distribution, and ori-
gin of many annual and perennial C4 grasses in Israel can
be found in Vogel et al. (1986). Both annual and perennial
grasses can be used for diverse green roofs with year-
round plant cover. Setaria viridis (L) PBeauv, Eragrostis
ballerii Daveau, and Cymbopogon parkeri Stapf are pre-
sented hereafter as examples of annual and perennial C4
grasses originating in the Mediterranean and Middle
61
Eastern regions. Setaria viridis is an example of a summer
annual C4 grass, which belongs to the NADP malic
enzyme malate formers (Vogel et al. 1986). It is character-
ized by fast life cycle of six to eight weeks, and its exten-
sive seed dormancy transcends its life cycle by more than
two times, which enhances its survival as a weed species
in crop fields. Currently, this species is being tested for its
suitability as a crop for bioenergy (Petti et al. 2013). Era-
grostis barrileri is another summer annual grass, which
belongs to the aspartate-forming NAD malic enzyme C4
grasses (Vogel et al. 1986). It is adapted to human-
induced disturbances and grows in the rangelands of the
Mediterranean and Saharo-Arabian phytogeographic
regions (Al Harthi et al. 2008). Cymbopogon parkeri is a
winter perennial C4 grass that is native to the Saharo-Ara-
bian phytogeograhic region and mostly occurs on rock
crevices (Vogel et al. 1986). The genus Cymbopogon is
known worldwide for its essential oil content (Avoseh
et al. 2015). Lemongrass (Cymbopogon citratus Stapf)
displays high productivity under dry conditions. The plant
is used as an antibacterial, antidiarrheal, and antioxidant
compound (Shah et al. 2011). Furthermore, lemongrass
oil can significantly reduce and sometimes completely
retard several postharvest fungal spores (Tzortzakis &
Economakis 2007). Many of the C4 grasses native to the
Mediterranean area are severely invasive, for example
Sorghum halepense (L), which is an aggressive weed spe-
cies worldwide (Follak & Essel 2012), or the summer
annual C4 grass Echinochloa colona (L), which is the
most common grass weed in tropical rice fields (Chauhan
& Johnson 2009). These species are strongly not recom-
mended for cultivation on green roofs outside their region
of origin.
Tidestromia oblongifolia, Standl. (Arizona honey-
sweet) is a perennial C4 dicot shrub that belongs to the
Amaranthaceae family. This summer-active species is
native to the Central American deserts and is highly
adapted to extremely arid environments (Bjorkman et al.
1972). T. oblongifolia has the highest photosynthetic opti-
mum among vascular plants (Berry & Bjorkman 1980).
Due to its low height yet high spatial propagation, its
growth pattern is associated with a high coverage level
that makes it attractive for rooftop gardens. In the domes-
tic context, the flowers are fragrant and attractive to bees,
butterflies, and birds.
Plants exhibiting CAM photosynthesis are extremely
tolerant to drought (Sayed 2001). By opening their sto-
mata to collect CO2 in the dark, storing it in the form of
an organic acid, and decarboxlyating it back during day-
light for the normal reduction cycle, CAM plants mini-
mize both transpiration losses and water uptake from the
soil (Cushman 2001, Ting 1985). Plants of the Sedum L.
genus, which belongs to the family of Crassulaceae, have
the ability to alternate between C3 and CAM pathways
(Borland & Griffiths 1990; Lee & Kim 1994; Werthmann
2007), which is beneficial on green roofs with unpredict-
able water supply. Further, Sedum species are character-
ized by a minimal root system (Van Woert, et al. 2005),
with high biological activity that keeps the soil tempera-
ture on the roof higher than that of the external air and
reduces the risk of freezing temperatures during winter
62 I. Dirks et al.
(Eumorfoponlou & Aravantinos 1998). Their tolerance to
extremely high and low temperatures makes them ideal
for green roofs in desert regions with high daily tempera-
ture amplitudes and minimum values below zero during
winter months. Plants of the Sedum genus comprise more
than 500 species worldwide, and they are currently the
most popular plants for use on green roofs (Klett et al.
2012; Van Woert et al. 2005; Werthmann 2007).
Dew and atmospheric water vapor as a potential water
source on green roofs
Extensive green roofs are characterized by shallow soils,
thereby requiring continuous watering of the plants in
order to prevent them from drying out. A long-term study,
in several sites in Germany, revealed that precipitation
had the largest impact on plant success and diversity on
green roofs, which indicates the importance of water
availability (K
ohler 2006). In the case of limited precipi-
tation and shallow soils, other water sources, such as fog,
dew, and high relative humidity, could play pivotal roles
for plant growth on green roofs. Fog arises when atmo-
spheric water concentration reaches saturation (Kidron
1999), whereas dew forms when the surface temperature
is equal to or drops below dewpoint temperature, and
water vapor condenses on cold surfaces (Agam & Berliner
2006). Dewfall rates on a plants canopy can reach
0.09 mm h1 (Garratt & Segal 1988). Higher altitudes and
topography with differences in temperature between day
and night promote atmospheric inversions, which often
are accompanied by fog and dew. When the surface tem-
perature is higher than the dew point temperature, water
vapor is absorbed until the moisture content of the absorb-
ing material is in equilibrium with the ambient air humid-
ity (Kuehn et al. 2004). In comparison to unpredictable
precipitation events, dew and high relative humidity can
be very reliable water sources. During 80% 90% of the
nights, the maximum relative humidity reached at least
80% during the rainless summer season in the Judean
Foothills and on the Carmel Ridge in Israel (Dirks et al.
2010).
The desert plants Salsola inermis Forssk. and Artemi-
sia sieberi Bess. receive around 50% of their total water
from dew alone rather than from unpredictable and insuf-
ficient local precipitation. Salsola inermis is an annual
plant that belongs to the Amaranthaceae family. Its distri-
bution is limited to saline and disturbed habitats in arid
and semiarid regions (Barazani & Golan-Goldhirsh 2009).
Artemisia sieberi is a perennial dwarf shrub of the com-
posite family that grows in arid and semiarid regions. It
belongs to the Irano-Turanian phytogeographic region
and has a wide distribution in the Mediterranean area, the
Middle East, and Central Asia (Mohamed et al. 2010).
The plant is heavily branched from the base and has small
silver and densely haired leaves that are replaced by small
scale summer leaves by the end of the rainy season. The
flowering season is autumn (Ghazanfar 2012).
Moisture uptake by these plants was positively corre-
lated with the dew accumulation on leaf surfaces (Hill
et al. 2015). Plants activate different techniques in order
to use water obtained from dew during the night or early
morning hours when they can benefit from this resource.
For example, funnel-shaped leaves channel the dew water
into the soil for root uptake. In the Negev Desert, a nega-
tive correlation between dew uptake and root depth was
found (Hill et al. 2015). Because of their abilities to use
dew as a water source for growth and photosynthesis, the
desert plants Salsola inermis and Artemisia sieberi would
be a good choice for cultivation on extensive green roofs
with shallow soils of less than 20 cm soil depth.
Halophytes and xerophytes impact productivity and
diversity, and improve visual quality
The tolerance of halophytes to osmotic stress, ionic spe-
cific stress, and nutritional imbalance derives from several
mechanisms, such as the production and control of osmo-
protectant molecules (Lefevre et al. 2009), the adjustment
of root architecture (Shelef et al. 2010) and function
(Rewald et al. 2013), succulence (Jennings 1968), salt
accumulation (Shelef et al. 2012), and more. Xerophytes
are halophytes that are adapted to extreme environments
(Manousaki & Kalogerakis 2011), and therefore, they
have a good record of suitability for green roofs. Salt
remediation is an additional benefit of xerophytes (Shelef
et al. 2012).
Several halophytes could display high performance
growth on the green roof environment. Lampranthus blan-
dus, a succulent shrub native to South Africa, is well
known for its tolerance to extreme conditions and its good
esthetic performance (Ben-Dov et al. 1993; Choukr-Allah
et al. 1996; Ben-Dov et al. 2001). Drosanthemum hispi-
dum (L.) Schwantes has minimum irrigation needs to
maintain its visual quality (Pittenger et al. 2001). The pre-
sented plants are well known for their persistence and
robustness, which together with their high visual quality
make them suitable candidates for green roofs.
Interactions between neighboring plants
Plant growth on green roofs is constrained by the bound-
aries of the roof area and the shallow soils that limit space
for root and foliar development. Adjacent plants may
form a dense environment with a high level of resource
competition due to mutual shading or competition
between root systems. If space is limited and plant density
is high, allelopathy might be highly relevant, as well as
any resource competition.
Allelopathy describes the inhibitory effect of a plant
on its surrounding environment and vegetation (Bhowmik
& Doll 1984; Xu et al. 2012). Secondary metabolites con-
stitute the most important group of allelochemicals (Belz
2007). They play both major defensive and attractive roles
in interactions between plants, as well as between plants
and herbivores, and between pathogens and pollinators
(Briskin 2000). Allelochemicals are produced in a variety
of plant organs, mainly in leaves, and, to a lesser degree,
in stems and roots. They are released by plants into the
atmosphere or soil by volatilization or leaching. Leaching
may also occur through the decomposition of plant resi-
dues and exudation from plant roots into the soil environ-
ment (Rachmilevitch et al. 2006a; Rachmilevitch et al.
 Israel Journal Of Ecology & Evolution
2006b; Blum 2011; Thorpe et al. 2011; Miranda et al.
2011; Weston & Mathesius 2013).
The desert shrubs Origanum dayi Post, Artemisia
judaica L, and Artemisia sieberi exhibited high allelo-
pathic effects in their natural habitats in the Negev Desert
(Friedjung et al. 2013). Origanum dayi Post is an endemic
subshrub found in the northern Negev and Judean Deserts
in Israel. Its principal habitats are soil pockets and crevi-
ces in smooth-faced limestone, flint outcrops and cliffs,
and adjacent dry river beds; it is locally abundant at alti-
tudes from 300 m below sea level to 600 m above sea
level (Danin 1983). Artemisia judaica L. is a perennial
shrub with small silvery leaves and bright yellow flowers
that appear between March and April. It grows in dry river
beds in the eastern Sahara, as well as in southwestern
regions of the Middle East. In Israel, A. judaica can be
found throughout the Arava Valley, between 340 m below
sea level and 500 m above sea level. A. sieberi was
already discussed.
Among the three desert shrubs, O. dayi, A. judaica and
A. sieberi, water deficit was the key driver for selective
metabolomic shifts (Friedjung et al. 2013). The release of
allelochemicals constrains the germination and growth of
adjacent plants. Regarding the allelopathic effects of the
abovementioned species, O. dayi showed comparatively
higher germination-inhibition effects, while A. judaica
displayed relatively higher growth-inhibition effects
(Friedjung et al. 2013). The essential oil of A. judaica can
be used for weed control (Dudai et al. 2003; Friedjung
et al. 2013). The constraint on the germination and growth
of adjacent plants results in subsequent effects on resource
availability and the level of competition on green roofs.
Regarding water availability, annuals, especially grasses
with strong tap roots, might be strong competitors (Vila &
Sardans 1999), and their limited growth can increase
resource availability and facilitate the growth of shrubs
and other herbaceous species.
Medicinal plants on green roofs
Medicinal desert plants, as a result of their exposure to
environmental stress, have developed unique survival sys-
tems based on secondary metabolites with remarkable
properties for human health (Harlev et al. 2012; Friedjung
et al. 2013). In response to drought stress and high light
intensity, plants close their stomata, thus constraining
CO2 uptake and suppressing the Calvin cycle. Hence, the
oversupply of NADPHC and HC pushes metabolic pro-
cesses towards the production of reduced compounds,
such as isoprenoids, phenols, and alkaloids (Selmar &
Kleinwachter 2013a; Selmar & Kleinwachter 2013b).
Many of these metabolites diminish oxidative processes
in the human body and impart antifungal, as well as anti-
bacterial, antiviral, anthelmintic, and antimutagenic capa-
bilities to the plant (Briskin 2000; Harlev et al. 2013).
The natural flora of Israel contains a wealth of plant
species with medicinal properties that have been used
since ancient times (Lev & Amar 2000). Desert plants
that are adapted to extreme environmental conditions and
that simultaneously have medicinal properties can
63
contribute to the achievement of ecologically and eco-
nomically sustainable gardening on green roofs. Second-
ary metabolites can be unique to specific species or
genera and are divided into three major groups based on
their chemical structure and biosynthetic origins: terpe-
noids, phenylpropanoids, and alkaloids. Terpenoids form
the largest class of plant metabolites, and their major
function in plants is chemical defense against insects and
environmental stress. Plant-derived terpenoids can be
involved in the repair of wounds and injuries (Bohlmann
& Keeling 2008) and can have beneficial therapeutic
effects against inflammatory diseases and cancer (Grass-
mann 2005; Salminen et al. 2008).
As a result of stress, the phenylpropanoid metabolism
in plants transforms phenylalanine into a variety of impor-
tant secondary products, including lignins, sinapate esters,
stilbenoids, and flavonoids (Vogt 2010). Phenylpropa-
noids can have anti-allergic, anti-inflammatory, antith-
rombotic, and anticancer effects on human health.
Flavonoids and stilbenoids have powerful antioxidant
properties, hence offering remarkable protection against
damage caused by free radicals (Korkina 2007; Ververidis
et al. 2007). Alkaloids are another major group of plant
secondary metabolites. They constitute a structurally
diverse group of over 12,000 cyclic nitrogen-containing
compounds that are found in over 20% of plant species. In
many cases, alkaloids directly interact with molecular tar-
gets within the nervous system, which act as feeding
deterrents and toxins to insects and other herbivores.
Many alkaloids are well known for their pharmacological
effects, including atropine, morphine, codeine, heroin,
nicotine, caffeine, cocaine, and quinine (Kennedy &
Wightman 2011).
Currently, natural products from medicinal plants
have an important function in medicine. Secondary
metabolites with a positive impact on human health were
detected in different medicinal desert plants.
Origanum dayi is known for its unique flavor and ther-
apeutic properties used to treat disorders of the digestive
and respiratory systems. Shoots of O. dayi contain essen-
tial oil constituting up to 2.8% of the fresh weight.
Approximately, 50 volatile constituents of the oil were
identified as belonging to the terpenoid group. The essen-
tial oil of O. dayi has significant economic potential for
the aromatherapeutic market, as it is the most expensive
essential oil produced from the Origanum genus (Dudai
et al. 2003). Artemisia judaica is strongly aromatic and
commonly used by the local population for various medic-
inal purposes, such as treating parasites in the digestive
system, gastrointestinal disorders, snake and scorpion
bites, ear infections, dysentery, coughing, and external
injuries. The upper parts of the plant contain essential oil
from the terpenoid and phenylpropanoid groups amount-
ing to 1.6% of the fresh weight (Putievsky et al. 1992).
Artemisia sieberi contains various secondary metabolites,
which include sesquiterpene, lactones, and flavonoids. Due
to its essential oil and other secondary metabolites, A.
sieberi is considered to produce several therapeutic effects
including: antioxidant activity, antivenom activity, antibac-
terial activity, antispasmodic activity, anthelmintic activity,
64 I. Dirks et al.
antileishmanial activity, neurological activities, hypoglyce-
mic activity, cytotoxicity, and gene induction (Mohamed
et al. 2010). Information about the concentration and total
amount of plant secondary products, in response to drought
stress, of various species and medicinal plants can be found
in Kleinwachter and Selmar (2015).
Establishment and maintenance of diverse green roofs
and ecosystem services under varying environmental
conditions
Green roofs can have a substantial impact on urban envi-
ronments. So far, green roof designers have mostly
focused on the energy consumption of buildings, air qual-
ity, and surface runoff in urban areas. These services have
mostly relied on plants of the genus Sedum.
The establishment and the maintenance of more diverse
green roofs should be based on both habitat features and
the inclusion of different life forms, such as perennials and
annuals. Habitat heterogeneity can be increased by incorpo-
rating physical habitat features, such as sun exposure, soil
depth, and different substrates (Willams et al. 2014).
Surfaces with high condensation, such as cobbles and
stones, can serve as a sink for nutrients in desert areas, due
to the fact that the concentration of nitrogen, phosphorus,
and potassium in dew is 1.4 3 times higher, compared to
that of precipitation (Kidron and Starinsky 2012).
In addition, perennial plants might contribute to small-
scale habitat heterogeneity by affecting physical habitat
features and micro-topography (Lundholm 2015). The use
of different life forms provides better microcosm func-
tioning and resistance to environmental stress, mainly due
to niche completion and facilitation (Lundholm et al.
2010). In this way, perennial plants can facilitate the per-
formance of neighboring plants, by providing a nursery
effect for other herbaceous species, when environmental
conditions are harsh (Butler & Orians 2011). Field
research on interactions between shrubs and annuals
reveals that this relation is still more complex and
depends, to a large extent, on water availability
(Thielb orger & Kadmon 2000; Friedjung 2013, Holzapfel
et al. 2006, Holzapfel & Mahall 1999). In Californias
Mojave Desert, an overall positive effect of shrubs on bio-
mass and the performance of annuals has been found,
whereas annuals had negative effects on shrubs, by means
of competition for water and other resources (Holzapfel &
Mahall 1999). Research from Israel provides evidence for
an increase in the positive effects of shrubs on annuals
with increasing rainfall, even though the magnitude of
this effect changed with the identity of the respective
annual plant. Explanations can be found in the intercep-
tion of rainfall by shrub canopies that were important in
limiting the emergence and reproduction of understory
plants during dry years (Thielb orger & Kadmon 2000).
Furthermore, desert shrubs increase their allelopathic
effects under drought, which constrain the germination
and growth of annuals (Friedjung 2013).
Mediterranean vegetation can be characterized by a
very high diversity of annuals (Caneva et al. 2013), but
these plants are rarely used on green roofs. Recently, a
broad study on the traits of annual plants indicated the
suitability of numerous species for the green roof environ-
ment by using functional traits (Van Mechelen et al.
2014a). Explanations for the underrepresentation of annu-
als on green roofs might include the fact that they are
absent during summer months, with a low contribution to
temperature insulation during this period (Van Mechelen
et al. 2014b). Seed dormancy might be considered as
another critical factor. However, a recent analysis of seed
traits, dormancy, and germination along a gradient from
the Mediterranean to the arid region in Israel revealed rel-
atively high germination rates of the dominant annual spe-
cies (50% 90%) and a significant increase in germination
rates with increasing aridity (Harel et al. 2011).
High levels of diversity could partly overcome the
issue of short aboveground visibility, provided that both
early and late season species are selected. Information
about the life forms of all presented species can be found
in Table 1.
Diverse green roofs have a better survival probability
and are more esthetically pleasing than species-poor green
roofs, even under dry conditions (Nagase & Dunnett
2010). This can have positive effects on life quality and
can increase the esthetic and economic value of buildings.
More diverse vegetation on green roofs might further sup-
port a range of local invertebrates with a subsequent posi-
tive effect on bird communities (Oberndorfer et al. 2007;
Madre et al. 2013; Madre et al. 2014).
Conclusion
Various plants native to the Negev Desert are adapted to
extreme heat and radiation, water stress, and salinity.
Thus, they could provide perennial cover and create an
esthetically pleasing environment. Furthermore, many
desert plants have medicinal properties that can contribute
to more diverse economic and cultural activities, making
city living more attractive. More diverse green roofs could
help to preserve and encourage the presence of inverte-
brates and birds in urban areas. To create and maintain
diverse green roofs, it is essential to consider the biotic
interactions between plants, especially competition for
resources and allelopathy, with respect to the fact that
they change with habitats and environmental conditions.
Disclosure statement
No potential conflict of interest was reported by the authors.
ORCID
Inga Dirks http://orcid.org/0000-0002-1362-9516
Amir Eppel http://orcid.org/0000-0001-7572-8424
Moses Kwame Aidoo http://orcid.org/0000-0003-0183-4771
Amnon Kochavi http://orcid.org/0000-0002-0238-711X
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