More About Coccoliths ...
Jeremy Young
Palaeontology Dept.
The Natural History Museum
London, SW7 5BD
Great Britain.
Email: j.young@nhm.ac.uk
Some more advanced discussion of coccolith shape and formation.
Chirality: one of the many remarkable features of the coccolith
structure is that they show pervasive and consistent chirality, or
handedness. Although there is a general radial symmetry to the structure in
detail almost every aspect shows also rotational symmetry. For instance the
distal shield elements are deflected clockwise when seen in distal view) and
the inner tube elements are deflected anti-clockwise. Again the proximal
shield elements are terminated by a pair of faces of the which the one on
the clockwise side is always longer. These chiral features are not always
perfectly developed but whenever they occur they always show the same
handedness - this has been tested by checking numerous specimens and
examples of reverse chirality have never been found. The chirality extends
back to the earliest growth stages and appears to be a product of the
nucleation stage.
Number of crystal units: whilst the structure of the crystal units is
extremely regular the actual number of crystal units is variable. Typically
an Ehux coccolith has about 30 crystal units - this can easily be
determined by counting the number of hammer head elements in the distal
shield. Small specimens, however, may have as few as 20 crystal units and
large specimens more than 40. As with the arrangement of coccoliths in the
coccosphere this appears to be a self-organising type of structure. There
is no precise coccolith blueprint, instead the crystal units nuclei are laid
down at more or less uniform spacing around the proto-coccolith ring so if
the ring is larger there are more crystal units.
Crystallographic orientation: calcite is a mineral with a precise
chemical composition (CaCO3) and atomic structure with its own internal
symmetry. Inorganic calcite crystals have shapes which reflect the internal
symmetry, typically rhombohedra or hexagonal prisms elongated along the
dominant c-axis of the atomic structure. In coccoliths the shape of the
individual crystal units is strongly modified but the c-axis orientation is
precisely related to their shape, specifically the c-axis is parallel to the
elongation of the hammer-like distal shield elements. We can conclude from
this that the initial nucleation stage of coccolith formation is
characterised by precise control of crystallographic orientation, as well as
the spacing and location of nuclei.
V&R model of nucleation: Ehux coccoliths despite their
complexity are actually somewhat simpler structures than many others in that
the entire structure is formed of only type of crystal-unit. In most other
coccoliths the proto-coccolith ring consists of two crystal unit types one
with approximately radially oriented calcite crystals (R-units), as in
Ehux and the other with approximately vertically directed c-axes
(V-units). These two crystal unit types can be traced back to the
proto-coccolith ring where the two alternate, V-R-V-R-V-R. Whilst only
R-units are seen in fully grown Ehux coccoliths study of
proto-coccolith rings shows that two types of crystal units occur in the
earliest growth stages; larger crystal nuclei which develop into the R-unit
crystals of the complete Ehux coccoliths and smaller crystal nuclei
which do not grow. We surmise that the smaller nuclei are relict V-units
which are superfluous to the final coccolith structure but which are still
nucleated since the nucleation stage is based on a conserved mechanism
which cannot readily be changed during evolution.
Biochemical control: ultimately biomineralization must be the product
of biochemical systems and we believe that with coccoliths we are close to
being able to determine and characterise them. This is a tantalising goal
since if we can understand how coccoliths, one of the most precisely
regulated biomineral structures known are formed then we will be a
significant step closer to understanding biominerization in general and to
being able to create biomineral like structures. It seems likely that two
separate system are involved.
1. Nucleation regulation: the evidence described above indicates that
the nucleation of calcite is very precisely controlled. A straightforward
hypothesis for explaining this is that an organic macromolecular template
exists with an array of binding sites which mirror the structure of a plane
through the calcite structure. On such a template Ca2+ and CO32- ions will
fit into the binding sites automatically producing a proto-calcite crystal.
Subsequent crystal growth can then occur inorganically. The alternation of
V and R nuclei seen in coccoliths has lead us to speculate (Young et al.
1992) that the macromolecular template is folded so that a single template
can produce the alternating nuclei with alternating vertical and radial
c-axes.
This template is entirely hypothetical but if it exists we would expect it
to be a protein directly controlled by RNA. Biochemistry research groups in
Japan and Europe are attempting to isolate this coccolith precursor template
molecule.
2. Growth regulation: an extremely complex acidic polysaccharide has
been identified by the Leiden research group (de Jong et al. 1984). This
has been shown to be closely associated with coccolith growth and to have a
high affinity for calcite. One likely hypothesis is that this molecule
binds to calcite and blocks further mineral growth on surfaces to which it
binds. This effect in combination with vesicle growth in particular
directions under cytoskeletal control may be sufficient to regulate crystal
growth.
Genetic control: the approach to understanding coccolith proto
described above has been to work down from observed coccolith structure
toward the biochemical systems controlling growth regulation. An
alternative approach is to attempt to identify and characterise the genes
responsible for regulating biominerization. This should be possible since
in culture strains often lose the ability to calcify, i.e. the genetic
systems which promote calcification are suppressed, so by comparing the RNA
produced by calcifying and non-calcifying Ehux strains the genes
regulating calcification should be identifiable. This type of enquiry is
being followed by a Dutch research group lead by Prof. Westbroek.
Our hope is that in this area as others the common focus on a single
species, Ehux, will enable us to address fundamental problems using a
wide range of different techniques and so develop detailed understanding of
one system as a model for wide application.
References
Ehux
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