by Michelle Farrell (@DrM_Farrell)
To around 20% of the UK
population, pollen is familiar as the cause of hay fever during the spring and
summer months. To a very small minority (ironically, many of them well acquainted
with the runny-nosed, itchy-eyed symptoms of hay fever themselves), pollen is
much more than an allergen. Pollen grains contain the male gametophyte of
seed-producing plants, and in order to increase their chances of reproductive
success, wind-pollinated plants produce pollen in vast quantities. The small
size of pollen grains (generally in the order of 20 to 40 microns, a micron
being one thousandth of a millimetre) means that when they are released by a
plant they become widely dispersed in the environment. As the plant has no
control over where its pollen grains end up, another part of the reproductive
strategy is that the grains have a very tough outer casing or exine,
made of a substance called sporopollenin, allowing them to survive in less than perfect conditions. This outer wall can be preserved in several
environments, particularly waterlogged ones, for tens of thousands of years.
This combination of pollen being produced in large quantities, wide dispersal in
the environment, excellent preservation under the right conditions, and the
fact that pollen grains can often be identified to family, genus and even
species level, is what makes them such a valuable tool for research.
|
Coring to retrieve pollen-bearing sediments from a bog
on Orkney, Scotland |
Pollen analysis is one of the most common methods used for investigating
past environments. Pollen is often preserved in waterlogged environments where
sediments accumulate, such as lakes and peat bogs. Cores of sediment can be
extracted from these locations, and sub-samples from various depths are then
subjected to a series of physical and chemical laboratory treatments which
remove the majority of the inorganic sediments and large organic debris,
leaving behind the fine organic fraction of the sediment. It is this fraction
that contains the pollen grains, as well as other tiny organic remains
including fern and fungal spores, other fungal remains such as hyphae, and
fragments of charcoal from either natural or anthropogenic fires. The study of
all these remains together is known as palynology, a term coined by the British
scientists Hyde and Williams in 1944 and derived from Greek words meaning ‘the
study of small particles sprinkled about’.
|
The stripes in this core segment indicate that the sediments
were deposited under different environmental conditions |
Once you have concentrated the fine
organic fraction of the sediment, the next stage is to identify the botanical
remains contained within it. Much as a botanist would use a key to help them
to identify plants out in the field, palynologists use keys to pollen and
spores to aid their identification of specimens under the microscope in the
lab. The identification and study of fungal remains is still a relatively new
technique, and to date no definitive key to these types of remains has been
published. Therefore I’ll focus here on the distinctive characteristics of
pollen grains that allow palynologists to distinguish between the different
taxa present in a sample. One of the most distinctive features of pollen grains
is their apertures. There are two types of aperture: pori (pores), roughly
spherical in shape, and colpi (furrows), which are elongated and have pointed
ends. Some grains have both colpi and pori
in the same apertures, and are known as colporate. The number and arrangement
of the apertures is also key in pollen identification. The number of apertures
is indicated by the use of the prefixes mono-, di-, tri-, tetra-, penta- and
hexa- before the terms porate, colpate and colporate. The prefix poly- is used
to denote the presence of more than six apertures. Usually the apertures are
arranged equidistantly around the equator of the pollen grain, and this is
indicated by the prefix zono-. Panto- is used when the apertures are scattered
all over the surface of the grain. Some examples of the way in which apertures
are used to identify pollen grains are shown below.
|
Betula (birch) pollen: trizonoporate,
with three pores arranged equidistantly
around the equator of the grain |
|
Fraxinus excelsior (common ash) pollen:
trizonocolpate, with three furrows arranged
equidistantly around the equator of the grain |
|
Rumex obtusifolius (broad-leaved dock):
tetrazonocolporate, with four apertures
made up of both colpi and pori |
|
Plantago lanceolata (ribwort plantain):
polypantoporate, with many pores scattered
all over the surface of the grain |
The pattern of
sculpturing found on the surface of the exine is another crucial factor in the
identification of pollen grains. Around fifteen different surface patterns have been
described, and two of the more distinctive patterns are shown in the images below.
|
Cirsium arvense (creeping thistle) pollen, a
good example of echinate surface sculpture |
|
Ulmus (elm) pollen, displaying rugulate
surface sculpturing |
Size
can also be important in distinguishing between pollen taxa, particularly
members of the grass family. The two images below show the difference in size
between a wild grass, Phragmites
australis (common reed) and a cultivated grass, Triticum aestivum (wheat).
|
Phragmites australis (common reed) pollen |
|
Triticum aestivum (wheat) pollen |
The identification and
recording of pollen grains from different depths within a sediment core can be
used to provide information on how the vegetation surrounding the core site
has changed over time. Sediment cores can often be accurately dated using
radiocarbon, allowing the changes in environment to be tied to a chronology.
Palynology has great potential for providing baseline data for the development
of conservation management strategies, and is also useful from an ecological
perspective as it can give insights into how plant communities have responded
to climate change in the past, thereby allowing predictions to be made about
how vegetation and ecosystems may be affected by future climate change.
One of my main
research interests is in the use of palynology as a tool to unpick the ways in which
humans interacted with their environments during the Holocene (the period since
the end of the last ice age, approximately 11,500 years ago, until the present
day). I intend to write more about the archaeological applications of
palynology in future posts, but as a taster, it is possible to determine when
people began farming in an area, what crops they were growing, where they
grazed their animals, whether they cleared woodland to create more land for
agriculture, and how they contributed to the development of cultural landscapes
such as heathlands. It is also possible to investigate the ways in which people
may have managed their environments and responded to climate change in the
past, for example managing heathland by deliberate burning in order to maintain
the quality of grazing. It may even be possible to detect woodland management
practices such as coppicing, and Jane Bunting will write about her work on this
in the next GEES-ology post.
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