Lichens
as Environmental Monitors
For those with a modicum of familiarity with lichens, it may come
as a bit of a surprise that they are widely used to monitor environmental
quality. Their use as so-called bio-indicators belies their well-earned
reputation for stolid tenacity under Stygian conditions. Lichens are globally
distributed with about 14,000 species that extend across a broad range of sizes
and shapes and inhabit every niche from the frozen polar extremities to the
equatorial tropics, favoring places where nothing else will grow; they are the
dominant form of vegetation for about 8 percent of the earth’s surface. The
seeming oxymoron of lichen sensitivity deserves some consideration that begins
with a few fundamentals of lichenology.
A lichen is "an association of a fungus and a photosynthetic
symbiont resulting in a stable vegetative body having a specific
structure" according to the definition accepted by the International
Association of Lichenologists. In other words, it is not really a singular
entity but a mutualistic association of two species from different kingdoms;
the fungus from Kingdom Fungi and the alga from Kingdom Plantae. Since fungi
are heterotrophic like animals, they must get their nutrition from an
autotrophic plant, the alga. The fungus provides structural stability, water
and minerals and the alga provides complex carbohydrates. Lichens have been
anthropomorphically cited as “fungi that discovered agriculture." The
basic functional structure of a lichen is relatively simple. The fungal portion, called the mycobiont,
constitutes the bulk of the vegetative body; the algal component,
or photobiont provides photosynthetic derived nutrients. This relationship is generally characterized
as benign mutualism, a type of symbiosis in which both constituents share the
benefits of the association. However, it is probably better to characterize the
relationship as at least partially parasitic, for in virtually every case the
fungus penetrates the alga and absorbs about half of its nutrients; the
survival of the lichen depends on the alga replenishing lost cells through
photosynthesis.
Lichens are sensitive to environmental pollutants for two very
fundamental reasons. First and foremost is the epiphytic nature of their physiology;
they ingest air and water directly from the atmosphere and do not therefore
benefit from the filtering and concomitant cleansing effects of an intermediary
such as soil. Air and water are exchanged over the entire external surface of
the fungal body, which is called a thallus.
Lichens have no natural means to retain water (lacking protective cells)
and are thus hydrated and dehydrated several times over the course of a
day. The end result is that lichens are
immediately and directly sensitive to any anomalous additives to the air and to
the water, concentrating trace contaminants with cyclic celerity. The second
reason for their sensitivity is their mutualistic association, which is a
somewhat tenuous balance of nutrition and resources. If an environmental
pollutant has a deleterious effect on either the alga or on the fungus, then
the union is threatened to a greater extent than would be the case with a
single organism. Either the alga or the fungus may be affected individually or
the dynamic of their resource sharing may be; in any of the three possible
impacts, lichen debility results. Lichens are therefore well suited natural
environmental monitors.
The
conflated composition of the lichen led to some understandable confusion in the
nascent sciences of the Age of Reason that transcended medieval superstitions.
The French botanist Joseph Tournefort first placed the lichens in a separate
genus of the
plant kingdom in the late 17th Century. Carolus Linnaeus, in
establishing the taxonomic rules of orders, families, genera and species,
classified them as algae. His student, Erik Archarius, who is known as the
father of lichenology, placed them with the fungi (then part of the Kingdom Plantae
in the Phylum Thallophyta). However, it was not until the middle of the 19th
Century that William Nylander, a Finnish botanist who had emigrated from
Helsinki to Paris, correctly identified the duality of the fungal-algal lichen.
He is also credited with discovering chemotaxonomy, using chemical reagents
such as calcium hypochlorite in the analysis of lichen speciation, and with
establishing a relationship between the lichen population and prevailing
environmental conditions, the latter published in the Bulletin of the Botanical
Society of France in 1866 as Les Lichens
du Jardin du Luxembourg. Nylander
noted that the lichens in the verdant Paris garden were notably richer in
quality and quantity than in any other part of the city. He may accordingly not
be mistakenly referred to as the father of lichen environmental monitoring.
In the
most generalized sense, lichens are divided into five categories according to
their basic morphology: crustose, foliose, fruticose, squalmulose and
leprose. Crustose lichens are the most familiar,
as they are the tightly adherent crust-like growths typically found on any
relatively old stone surface like a grave marker. Foliose and fruticose are the
lichens that look like little leaves (folium
is the Latin word for leaf) or little ramified shrubs (frutex is the Latin word for shrub and has nothing to do with
fruit). The last two are essentially variants of the crustose variety;
squamulose lichens are crust-like with upturned scales (a squama is a scale)
and leprose lichens are crust-like with a loose powdery surface. On a
macroscopic perspective, the environmental health of an ecosystem can be
ascertained according to the types and distribution of
lichens; their absence in an otherwise appropriate location indicating that air
quality is exceptionally poor. Only crustose lichens are found in the most polluted
areas; as air quality improves, the more complex lichen types such as
squamulose and leprose become apparent. At the other end of the scale,
fruticose and foliose lichen are indicators of clean air. In our area, the
extensive colonies of the leafy foliose Rock Tripe (Umbilicaria mammulata) and the branchedfruticose Reindeer Lichen (Cladina rangiferina) at the upper
elevations are indicative of salubrity. The structure is evident in the
photographs of each above.
The practical use of lichens as environmental indicators is more specialized
in that it depends on the reaction of an individual species of lichen to a
specific pollutant and on the absorption of different air pollutants by the lichen
as an inherent consequence of the photosynthetic respiration process. The primary pollutant of concern is sulphur
dioxide (SO2), a strongly polar molecule that is introduced into the
atmosphere primarily as a by-product of the combustion of fossil fuels which frequently
have sulphureus composition. Sulphur dioxide adheres readily to the external
surfaces which results in a reduction in respiration and concomitant
photosynthesis of the algal component on which the lichen relies for sustaining
energy; it is
the primary agent for lichen mortality, particularly those of fruticose
structure due to their larger surface areas. Natural background SO2
ranges over one order of magnitude (0.28 to 2.8 milligrams per cubic meter –
mg/m3) whereas excessive combustion of high sulphur fuels can raise
this up to about 200 mg/m3; lichens are essentially exterminated
above levels of 60 mg/m3. The second major lichenous pollutant is
the result of the hydrolytic reaction of sulphur dioxide to sulphuric acid (H2SO4)
and of nitrogen oxide (NO2), also a product of fuel combustion, to
nitric acid (HNO3). The end result is acid rain, defined as
atmospheric water (rain or snow) with a PH of less than 5.6 (PH stands for pouvoir hydrogčn which
is French for power of hydrogen and is a measure of acidity; 7 is neutral and
<7 is acidic). Lichens subject to a PH of less than 2.5 exhibit reduced photosynthesis
and a decrease in weight. The third and final pollutant category affecting
lichen population diversity and mortality are metals in their ionic state.
Silver (Ag+), mercury (Hg+) and copper (Cu+)
are noted for their lethal toxicity while lead (Pb+), zinc (Zn+2)
and nickel (Ni+2) are of intermediate toxicity.
Lichen environmental monitoring has come a long way since Nylander
perambulated the pathways of Parisian parks and observed that they were more
lichenous than their adjacent urban environs. The first extension of the general
view of lichens as indicators was undertaken by the Swedish botanist Rutger
Sernander, who systematized Nylander’s lichen and non-lichen dichotomy into a
tripartite of a “lichen desert” zone, a transition or struggle zone, and a
normal or lichen growth zone which he applied to the city of Stockholm in the
1920’s; detailed lichen maps of Helsinki and Oslo were diagrammed in the
1930’s. The increased use of fossil fuel
that ultimately resulted in the coining of the word smog from smoke and fog to
describe the atmospheric effects incubated resurgence in monitoring with
lichens in the 1970’s. A ten zone system based on lichen sensitivity was
devised in the U. K. by David Hawksworth and Francis Rose to map the extent of
sulphur dioxide damage in Ireland, Wales and England. At about the same time,
the Canadians Fabius LeBlanc and Jacques DeSloover developed the Index of
Atmospheric Purity (IAP), a quantitative score based
on the relative plenitude of various lichen species, and used it to map
Montreal. More recently, the availability of sophisticated laboratory protocols
such as atomic absorption spectrophotometry and X-ray fluorescence spectrometry
has resulted in the capability to monitor elemental atoms that comprise the
lichen thallus. The analysis generally includes sulphur, nitrogen and fluorine
in addition to a wide range of metals from copper to zinc including lead and
chromium. For example, in 1993, the U. S. Forest Service in conjunction with
the Environmental Protection Agency initiated a lichen indicator section in the
extant Environmental Monitoring and Assessment Program/Forest Health Monitoring
(EMAP/FHM) Program at a number of test sites. The assessment consists of the
determination of 27 different contaminants in 10 different species of lichen
and includes a complete survey of epiphytic lichens in selected areas. Lichen
analysis was used to determine the extent of radioactive contamination
following the reactor accident at Chernobyl near Kiev in Ukraine.
So why do we use lichens to conduct atmospheric monitoring? It’s really
quite simple - because they do it for us. Were it not for lichens, each
sampling location would need to have an air monitor with sophisticated
electronics necessary for the requisite accuracy and precision in addition to a
power supply and a suction device to siphon air across a collection membrane of
some sort. This would be prohibitively expensive – and unnecessary. The extent to which lichens provide effective
and affordable environmental monitoring is manifest in the practice of
transplanting lichens into areas lacking sufficient populations to function as
natural air monitors. The bottom line is that lichens instantiate environmental
measurement systems.