Coral Coloration: Fluorescence: Part 1

IntroPhoto.jpg

An unidentified Hawaiian zoanthid fluoresces nicely under
blue light.

Science is a journey full of
surprises and the story behind the current interest in coral
fluorescence and pigmentation is no less full of twists and
turns. This tale is one of the reef-keeping hobby in Russia and
Japan, a couple of fortuitous laboratory accidents, a revolution
in biomedical research, revelations and fortunes made, discovery
of a new symbiosis, as well as the small contributions of reef
hobbyists from other parts of the globe. While coral coloration
is today less likely to be the subject of intense interest (and
debate) within the hobby than it was just 5 years ago, the
situation is much different in peer-reviewed literature, with
researchers producing a staggering amount of information on
marine invertebrate fluorescent and non-fluorescent pigments. It
is estimated that 26,000 papers mentioning fluorescent proteins
will be in print by the end of 2006. Not all of these –
few, in fact – are of much interest to the advanced
aquarist, however, some offer glimpses of those conditions
required to activate, or inactivate, the vivid coloration we
sometimes see in our captive reefs. Of course, explaining pretty
colors in an aquarium (or reef) is not usually researchers’
main goals. The revolutionary use of certain proteins as
molecular markers (or ‘highlighters’) in the
biomedical field has driven the identification, refinement and
genetic modification of many ‘wild type’ fluorescent
proteins. Other researchers have studied coral pigments in order
to remotely sense (by aircraft or satellite) the light reflected
and fluoresced by natural reefs. In this manner, vast tracts of
reefs can be examined in relatively little time and an assessment
of reef health can be made.

Why should reef hobbyists concern themselves with
fluorescence? Isn’t it generally accepted that light energy
is a prime requirement for expression of coloration? What if
could understand and anticipate responses of a pigment to
different visible light bandwidths, ultraviolet radiation or
other stimuli? What of the effect of nutrients and supplement
additions (such as iodine)? Not all pigments are fluorescent but
all are selective in the wavelengths they absorb or reflect. With
the recent introduction of relatively inexpensive spectrometers
(such as those from Ocean Optics or Edmund Optics) more hobbyists
can gain access to these powerful tools (for a review of the
Ocean Optics spectrometer, see Riddle, 2006). An understanding of
fluorescent proteins (at least their excitation and emission
peaks – see definitions below) is essential for
understanding the apparent color of corals, since fluorescence
can influence the color perceived by the observer. Further,
understanding of fluorescence and its effects is essential when
we examine reflected light later in this series. Realizing the
potential reasons for color transitions of our captive corals
might enable us to see these shifts as environmental indicators,
good or bad. But, perhaps most importantly to us as hobbyists,
manipulating the artificial conditions with aquaria might allow
us to alter pigmentation within certain invertebrates and create
animals with ‘customized’ coloration.

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This modest series of articles will review some recent
conclusions drawn from experiments from the world’s leading
authorities on fluorescent and non-fluorescent pigments. When
possible, recommendations of areas for further investigation by
hobbyists will be suggested. Observations by hobbyists can be of
great use. It is difficult to ignore mountains of anecdotal
evidence – it has been generally accepted within the hobby
that light is sometimes a critical factor in promotion of some
coral coloration.

In 1995, hobbyist (and scientist) Craig Bingman was perhaps
the first to recognize that the rainbow of colors in marine
invertebrates shared a common ancestor in a green fluorescent
protein; an addendum followed in 1999. All observations have not
proven prophetic – it was once very popular within the
hobby to associate all coral coloration with pigments found
within zooxanthellae. While it is true that light absorbed by the
photopigments of symbionts does play an important part in the
overall coloration perceived by the human eye, it is
generally a mistake to ascribe fluorescence to chlorophyll
and a definite error to link it to peridinin, beta-carotene and
some other photopigments (the role these pigments play in
determining the apparent coral color will be examined in a
separate article. Phycoerythrin is a notable exception as it can
lend an orange fluorescent color to corals under proper
conditions). Articles that state differently-colored
zooxanthellae are responsible for fluorescence are misleading. In
the same vein, coloration was wrongly attributed to natural
sunscreens that absorb and mitigate the effects of damaging UV
radiation. By extension, the idea was formed within the hobby
that ultraviolet energy is necessary for colorful corals (we know
now that these sunscreens – MAAs for mycosporine-like amino
acids – are colorless and do not lend color. We’ll
examine in this series the recent evidence for the necessity of
UV in color shifts and relate this to aquarium husbandry
practices).

Peer-reviewed publications have not always been correct
either. For instance, Kennedy (1979) described colorful
invertebrate pigments as porphyrins (naturally abundant in plant
and animal tissues. Porphyrin is a Greek word meaning
‘purple’ – they are chromatic pigments).
Kennedy reviewed early work by Moseley (1877) which describes the
red pigment found in Discosoma, Actinia,
Flabellum, Stephanophyllia and Fungia as
‘polyperythrin’ with excitation wavelengths in the
yellow-green portion of the spectrum.

Red fluorescence found in stony corals Montastraea
cavernosa
and Mussa angulosa tempted Kennedy to
describe them as porphyrins as well. Even with missteps the
pieces of the coloration puzzle are slowly but surely falling
into place.

For general interest articles examining coral fluorescence in
aquaria, see works of Riddle and Amussen (1998), Delbeek (2003),
Calfo (2005), Blundell (2005), Finét and Lesage (2005) and
Credabel (2006). Interesting comments concerning a few specific
fluorescent pigments can be found in Delbeek and Sprung, 2005, as
well as in Tyree (1998). Charles Mazel maintains a fascinating
website on marine fluorescence at
www.nightsea.com.

My fascination with coral pigments began when I saw a
blue-tipped Acropora (formosa?) specimen. This was
in the early 1990’s, and many hobbyists (including myself)
were of the opinion that it was practically impossible to
artificially over-illuminate an aquarium containing stony corals
such as Acropora. So, I was very curious why a coral would
reflect light energy necessary for photosynthesis (the coloration
did not appear to be fluorescent). I did not know it at the time,
but this simple observation would be a turning point in my life.
I’m still unsure of the reason why that Acropora
fragment reflected blue wavelengths, but my fascination would
become a passion in the quest for answers about coelenterate
coloration.

The first in this series will present information on a few of
the fluorescent proteins found in marine invertebrates.
Generally, fluorescent proteins are best viewed under relatively
narrow bandwidths, such as those produced by actinic and black
lights. However the mere categorization of
‘fluorescent’ and ‘non-fluorescent’
proteins presents difficulties – a protein can be
fluorescent in a technical sense but the ‘glow’ may
be of low intensity and invisible to the unaided eye. In
addition, environmental conditions (such as light quality and/or
quantity or chemical changes) may trigger a mutation of a
non-fluorescent protein to a fluorescent species or vice versa.
This leads us to a point where some definitions are in order
before we continue.

Fluorescence. Fluorescence is a phenomenon in which a
material absorbs light of one color (wavelength) and emits it at
a different color (wavelength). Absorption occurs when an
incoming photon (light particle) causes an electron to move from
a stable ground state to a higher energy, unstable excited state.
One of the ways for the excited electron to return to the ground
state is to ‘jump’ back down, emitting a photon of light. There
is always some energy lost to heat in the process, so the emitted
photon has less energy than the original photon. The energy of a
photon is related to its wavelength, which we perceive as color
– higher energy corresponds to shorter wavelengths, lower
energy to longer wavelengths. This shift in color was noted by
the brilliant, if slightly off-beat, Sir George Stokes, who first
described fluorescence in 1852.

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See this
informative web site
for further information.

Emission spectrum. The emission spectrum is a
measurement of the emitted energy as a function of wavelength. It
is generally presented as a graph (see Figure 1). The spectrum
will have a maximum but may also have secondary peaks, called
‘shoulders’. In text the spectrum is often described by
the wavelength of the emission peak. Some emission spectra cover
a broad range of wavelengths, while others are quite sharp. This
is described by the Full Width at Half Maximum (FWHM), the
spectral width in nanometers at the level that is 50% of the
value at the peak. The emission spectrum can be measured with a
basic spectrometer like the units from Ocean Optics.

Excitation spectrum. The excitation spectrum is a
measurement of the relative ability of different wavelengths of
light to stimulate (excite) the fluorescence (see Figure 1). Like
the emission spectrum, the excitation spectrum will have a
maximum but may also have secondary peaks. The excitation
spectrum tells you what wavelengths of light will be good at
making that particular substance fluoresce. Unlike the emission
spectrum, the excitation spectrum is difficult to measure without
specialized, expensive instrumentation. To make the measurement
you have to vary the wavelength of the incident light in very
fine increments and measure the fluorescence response at each
one.

Fluorescence lifetime. Fluorescence is short lived
– once the excitation wavelengths are absent, fluorescence
decays in a fraction of a second (billionths of a second), and
the phenomenon ends.

Stokes shift. The Stokes shift describes the difference
(in nanometers – nm) between the maximum excitation and maximum
emission wavelengths. Stokes shifts of coral pigments can range
from just a few nanometers to ~180nm and perhaps more.

Quantum Yield. The ratio of photons emitted through
fluorescence to photons absorbed by a pigment is called the
Fluorescence Quantum Yield. The higher the fluorescent quantum
yield, the more efficient a pigment is at fluorescing absorbed
light. Quantum yields and relative brightness vary significantly
among reports, and could be due to many factors including protein
purity and concentration, maturation conditions, protein source
and so on (Terskikh et al., 2002).

Relative Brightness. Relative brightness is the
comparison of emitted fluorescence to that of a standard (such as
the fluorescent pigment from the jellyfish Aequorea, or a
fluorescent dye). If the pigment’s ‘glow’ is
less intense than that of the reference, the relative brightness
is less <1. It is possible for the relative brightness to
exceed that of the standard in which case, of course, the
relative brightness will exceed 1.

Fluorescence Coupling and Förster Resonance
Energy Transfer (FRET).
Fluorescence Coupling occurs when a
pigment’s emission overlaps the excitation spectrum of
another pigment and energy is transferred between them via a
radiative mechanism (that is, a photon is emitted from one
pigment and absorbed by the other – this is a very low
probability event). In this manner, it is possible that
sequential transformation of wavelengths could occur. Coupling is
possible between coral fluorescent pigments (FPs). Förster
Resonance Energy Transfer (FRET, sometimes called Fluorescent
Resonance Energy Transfer) is a non-radiative,
‘vibrational’ energy transfer between intimate donor
and acceptor pigments. FRET is maximal when the distance between
pigment granules in corals is <10µm (Salih et al.,
2000). When it does occur in nature, energy transfer by FRET can
be very efficient.

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Photoconversion. Photoconversion is a change in
absorption and emission properties of a fluorescent protein
caused by incident light energy. Ultraviolet, violet, blue and
green bandwidths are known to cause photoconversions. At least
two sorts of photoconversion are known to occur in anthozoan
pigments. These are known as ‘reddening’ and
‘photobleaching.’ Generally ‘reddening’
describes a process by which green fluorescent compounds are
converted to those exhibiting red fluorescence.
Photobleaching usually describes a change from green
fluorescence to no fluorescence at all. Photoconversion is not
responsible for all ‘reddening’. Some color shifts
are due to:

Chemical oxidation. Oxygen plays a role in the
maturation (colorless to green color shift) in the green
fluorescent protein (GFP) of Aequorea victoria. It
also plays an essential part in ‘reddening’ of
fluorescent proteins in the false coral Discosoma, and
‘greening’ of a purple-red chromoprotein found in
Goniopora tenuidens. We’ll discuss photoconversion
in much more detail later in this series.

Photoactivatable Fluorescent Proteins (PAFPs). Some
proteins of interest to aquarists can convert from a
non-fluorescent state to one of fluorescence given the proper
environmental triggers. Visible light (of various wavelengths and
intensities) are required for photoactivation of these
‘chameleons’. Ultraviolet radiation can also activate
certain pigments (more on this subject later in this series
– don’t pull those UV shields yet!).

Figure1.JPG

Figure 1. An idealized chart demonstrating the concepts of
fluorescence excitation and emission spectra, as well as the
Stokes shift.

Figure2.JPG

Figure 2. Normalized excitation spectra of various
fluorescent pigments found in anthozoans. Note the distinct
double or triple ‘humps’ of Pigments 538, 575 and
583 – the lesser peaks are called
‘shoulders’. Spectra from various references.

Pigment Characterization

Researchers have variously reported fluorescent proteins by
color groupings (cyan-green-yellow-red), clade, numerical
emission wavelength, or as overt and covert fluorescence.
Fluorescence can play a major role in the appearance of an
organism (such as a coral illuminated by a black light or
spectrum weighted in the appropriate excitation wavelengths as
found in ‘actinic’ fluorescent lamps or high-Kelvin
metal halide lamps). Fluorescence can also play a part in the
perceived color of a coral in daylight – strong
fluorescence can ‘mix’ with the light reflected from
a coral (This mixing of fluorescence and reflected light will be
examined in a separate article). Not all coral pigments are
fluorescent. These definitions will be used in this series of
articles:

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Chromoproteins. For our purposes, chromoproteins will
describe the set of anthozoan pigments that do not visibly
fluoresce (if at all). This includes the non-fluorescent pink and
blue coral pigments (collectively called
‘pocilloporans’, Dove et al., 1995) and purple/red
pigments found in some corals and anemones. ‘CP’ is
the abbreviation for chromoprotein.

Fluorescent proteins can be subdivided into two
categories:

Photopigments are organic compounds that harvest light
energy for use in photosynthesis. Also called
‘accessory’ or ‘antennae’ pigments, the
fluorescent substances of most interest to us in the discussion
of coral fluorescence include chlorophyll a and
phycoerythrin.

Green Fluorescent Proteins and homologues. Green
fluorescent proteins (GFPs) and GFP-like proteins are those
fluorescent pigments not intimately associated with
photosynthesis (coral fluorescence is not due to
the color of zooxanthellae!). Fluorescent proteins are variously
colored cyan (CyFP), green (GFP), yellow (YFP) or red (RFP).
These may have some role in photoprotection of the coral host
and/or its symbiotic zooxanthellae. There are arguments for and
against this line of reasoning. This will be discussed later in
this series of articles.

‘Kindling’ Proteins (KPs) are sometimes
referred to as Kindling Fluorescent Proteins (KFPs) and do
not fall neatly into a fluorescent/non-fluorescent category as
they can be transiently fluorescent. The transformation of a
non-fluorescent protein to one capable of fluorescence is known
as ‘kindling.’ Light (quality- often green
wavelengths – and quantity or intensity) is often the environment
trigger for kindling. See this
site
for an interesting
tutorial:

Categorization of Coral Pigments

Various texts categorize visible radiation into divisions of
perceived color and associated wavelengths. These categorizations
do not always agree. I have unilaterally chosen a set of colors
and associated wavelengths from one of my fossilized college text
books and will list pigments accordingly (these are my articles
– I can do anything I want!).

There is yet one more definition – the method of
determining the shorthand for pigments. Again, there is no
universal agreement. Matz et al., 1999 and Labas et al., 2002 use
the genus name followed by a pigment number (I assume this is the
order in which the pigments were identified, such as
Zoanthus 1 or Discosoma 3). Matz et al. (1999) list
two pigments from Zoanthus, and common sense dictates that
we call them Zoanthus 1 and Zoanthus 2, yet Labas
et al. (2002) list a pigment called Zoanthus 2 that has a
different excitation and emission than either of the pigments
listed in the Matz paper. There must, hopefully, be a better
method. Some papers used abbreviations such as
‘mcavFP580’ (‘mcav’ means the pigment was
isolated from the stony coral Montastraea cavernosa, while
‘FP’ stands for ‘fluorescent protein’ and
the numbers describes the wavelength of the maximum fluorescent
emission). This is perhaps unwieldy for our purposes, and I for
several reasons have chosen to use the early (and shorter)
abbreviations such as P-575

(‘P’ is for ‘pigment’ and
‘575’ is the wavelength of maximum fluorescence
emission). There may be several coral genera containing a pigment
with the same fluorescent emission, and the same pigment may
exhibit small shifts in its exact peak location. The answer seems
to list each pigment, by emission, and list the host animal in a
separate column.

For non-fluorescent pigments (chromoproteins, or CPs)
the wavelength number stands for the maximum absorption. Bear in
mind that it is entirely possible for a single specimen to
contain multiple pigments (whether they are fluorescent, or not).
See Figures 3 and 4.

It is an important concept but most hobbyists are well aware
of this fact – this phenomenon is apparent when specimens
are viewed under ‘actinic’ fluorescent lamps.

Figure3.JPG

Figure 3. Though gaudily colored, the purple coloration of
this Acropora specimen is due to the presence of a
chromoprotein, and is not visibly fluorescent. The coral does,
however, contain a fluorescent pigment. See Figure 4.

Figure4.JPG

Figure 4. A fragment of the coral pictured in Figure 3, but
viewed under a black light emitting a UV-A radiation at a
maximum of 365nm. This green fluorescent pigment hides the
normal purple coloration of the animal. This Acropora
specimen has at least one fluorescent pigment and at least one
non-fluorescent chromoprotein.

Fluorescent and reflective pigments found in coral tissues are
generally water-soluble. Loss of pigments in nature or in an
aquarium would be expected to be quite low; however, corals
contribute fluorescent compounds to the water column (Gentien,
1981). This scientist demonstrated that amounts of fluorescent
compounds increased in prefiltered seawater when Acropora
formosa
, A. cerealis, A. hyacinthus, Stylophora
pistillata
, Favia favus, Favites flexuosa and
Leptoria phyrgia specimens were incubated.

Koh (1997) found that the non-photosynthetic coral
Tubastraea faulkneri secreted bioactive compounds to the
water column, and some of these compounds could be visualized
through fluorescence on developed thin-layer chromatographs.
Allelopathic intra-actions and interactions of coral color morphs
will be discussed briefly later in this series.

Phylogenetic Classifications

Though still in its infancy, computer software can sort (for
our purposes) zooxanthellae and even pigments into
‘clades.’ At present, at least four pigment clades
are known (A-D), which contain differing variations of cyan,
green, yellow and red fluorescent proteins plus non-fluorescent
chromoproteins. Figure 5 is one interpretation of pigments’
phylogenetic tree.

Software of this type is considered by some as subject to
fault, but it is an interesting concept.

Figure5.JPG

Figure 5. One interpretation of anthozoan colors’
phylogenetic tree (After Shagin et al., 2004). Clade
‘A’ GFP (found in some anemones) will fluoresce
bright green during its transition from green to red. Clade B
GFP demonstrates far less fluorescence than Clade
‘A’ during transition. Clade C GFPs from zoanthids
are like Clade ‘A’ in that they strongly fluoresce.
Clade D (mostly in stony corals) requires ultraviolet radiation
or violet/blue light to mature (Labas, 2002). Note: Mazel,
1995, describes a yellow fluorescent protein in the
scleractinian Agaricia.

A Listing of Fluorescence Proteins

Table 1 is a compilation of over 90 fluorescent proteins found
in various invertebrate species (non-fluorescent chromoproteins
will be listed under separate cover). I have color-coded the
appropriate fields to indicate approximate color of the
wavelength listed. The column on the left lists the
pigment’s name. Next is the maximum emission column
followed by two additional columns listing second and third
emissions (shoulders), if present. The excitation columns follow.
(Note: Observed excitation and emission spectra can vary between
instruments). Finally, the last two columns list the invertebrate
host (in alphabetical order) and then a reference listing for
those interested in further reading. I have used the genus and
species identifications listed within various papers. Coral
taxonomy is in a constant state of flux and I am not (nor do I
have the time to be) qualified to revise identifications. This
listing is as complete as I can make it with the references
available to me. A fluorescent emission peaking at a certain
wavelength does not necessarily mean that the animal host will
appear the color that the wavelength indicates (individual coral
colonies can contain ‘over 10,’ and possibly more,
fluorescent pigments, Salih et al., 2004). Perceived color is a
result of many factors.

Table 1. A compilation of anthozoan pigments reviewed in this series (listed by coral species).
PigmentEmission23Excitation23Found in:Reference
P-445445**340**Acropora asperaDove et al., 2001
P-490490514*480501*Acropora asperaPapina et al., 2002
P-500500**480**Acropora asperaDove et al., 2001
P-514514490*501480*Acropora asperaPapina et al., 2002
P-630630**576**Acropora asperaDove et al., 2001
P-480480510*481**Acropora aspera (green band)*Papina et al., 2002
P-476476510575500475*Acropora aspera (orange band I)*Papina et al., 2002
P-478478510575501475*Acropora aspera (orange band II)*Papina et al., 2002
P-487487517****Acropora cervicornisMazel,1995
P-518518*****Acropora cytheria @ Waikiki AquariumHochberg et al., 2004
P-490490**425**Acropora digitiferaDove etal., 2001
P-495495*****Acropora digitiferaGilmore et al., 2003
P-518518*****Acropora digitiferaHochberg et al., 2004
P-590590**570**Acropora digitiferaDove et al., 2001
P-485485**420**Acropora horridaDove et al., 2001
P-400400**345**Acropora horridaDove et al., 2001
P-625625**575**Acropora horridaDove et al., 2001
P-490490**405**Acropora milleporaCox and Salih, 2005
P-504504**405**Acropora milleporaCox and Salih, 2005
P-593593**405****Acropora milleporaCox and Salih, 2005
P-409409*****Acropora nastuaGilmore et al., 2003
P-482482515*451430*Acropora nastuaPapina et al., 2002
P-483483**427451*Acropora nastua (green band)*Papina et al., 2002
P-486486**384**Acropora nobilisSalih et al., 2000
P-484484515*450502*Acropora secalePapina et al., 2002
P-482482515*452500425Acropora secale (green band)*Papina et al., 2002
P-495495**472**Acropora sp.Karasawa et al., 2003
P-485485517555465505*Acropora tenuisPapina et al., 2002
P-517517555485505465*Acropora tenuisPapina et al., 2002
P-480480515*470504*Acropora tenuis (green band)*Papina et al., 2002
P-509509**397**Aequoria victoriaTsien, 1998
P-508-620508-620*****Agaricia agaricites @ 60mVermeij et al., 2002
P-565565**490**Agaricia humilisMazel et al., 2003
P-486486**426**Agaricia sp.Mazel, 1997
P-497497527****Agaricia sp.Mazel, 1995
P-513513545490***Agaricia sp.Mazel, 1995
P-515515*****Agaricia sp.Mazel, 1995
P-557557600545***Agaricia sp. Mazel, 1995
P-542542*****Agaricia undata @ 40mVermeij et al., 2002
P-486486**458**Anemonia majanoMatz et al., 1999
P-499499**480403278Anemonia sculataWiedenmann et al., 2002
P-595595**574**Anemonia sculataWiedenmann et al., 2000
P-522522**499***Anemonia sculata var. rufescensWiedenmann et al., 2000
CP-562***562**Anemonia sulcata, immature P-595Wiedenmann et al., 2002
P-565565**548**Cerianthus sp.Ip et al., 2004
P-685685**Violet/ BlueRed*Chlorophyll – common to healthy hermatypic coralsMultiple references
P-484484**456**Clavularia sp.Matz et al., 1999
P-515515*****Colpophyllia natansFux and Mazel, unpublished
P-496496**399482*Condylactis giganteaLabas et al., 2002
P-497497527****Condylactis giganteaMazel, 1995
P-508508**494**DendronephthyaLabas et al., 2002
P-575575**557**DendronephthyaPakhomov et al., 2004
P-486486**~448**Diploria labyrinthiformisFux and Mazel, unpublished
P-500500**475**Discosoma sp.Gross et al., 2000
P-583583**558530487Discosoma sp.Matz et al., 1999; Baird et al., 2000
P-593593**558**Discosoma sp. 2Fradkov et al., 2000
P-512512**503**Discosoma sp. 3Labas et al., 2002
P-593593**573**Discosoma sp. 3Labas et al., 2002
P-483483**443**Discosoma striataMatz et al., 1999
P-611611**559**Entacmaea quadricolorWiedenmann et al., 2002
P-487487517****Eusmilia fastigataMazel, 1995
P-518518**503**Family PectiniidaeAndo et al., 2004
P-517517**507**Favia favusTsutsui et al., 2005
P-593593**583**Favia favusTsutsui et al., 2005
P-507507536575***Favia fragumMazel, 1995
P-561561**548**Fungia concinnaKarasawa et al., 2004
P-505505**492**Galaxea fascicularisKarasawa et al., 2003
P-480480*****Goniopora tenuidensSalih et al., 2004
P-494494*****Goniopora tenuidensSalih et al., 2004
P-509509*****Goniopora tenuidensSalih et al., 2004
P-520520**488**Goniopora tenuidensSalih et al., 1999
P-582582*****Goniopora tenuidensSalih et al., 2004
P-500500**405481*Heteractis crispiaLabas et al., 2002
P-510510**490420*Heteractis magnificaTu et al., 2003
P-446(?)446**380**Leptoseris fragilisSchlichter et al., 1985
P-516516*****Lobophyllia hemprichiiNienhaus et al., 2005
P-483483**572454510Lobophyllia hemprichii (red)Salih et al., 2004
P-515515*****Lobophyllia hemprichii (red)Salih et al., 2004
P-519519*****Lobophyllia hemprichii (red)Salih et al., 2004
P-580580*****Lobophyllia hemprichii (red)Salih et al., 2004
P-581581*****Lobophyllia hemprichii (red)Salih et al., 2004
P-519-557519-557590****Madracis carmabiVermeij et al., 2002
P-520-555520-555590****Madracis carmabiVermeij et al., 2002
P-522-623522-623*****Madracis formosa @ 40mVermeij et al., 2002
P-534534575****Madracis formosa @ 60mVermeij et al., 2002
P-532532590****Madracis pharensis (brown tissue @ 10m)Vermeij et al., 2002
P-533533590****Madracis pharensis (brown tissue @ 20m)Vermeij et al., 2002
P-535535587****Madracis pharensis (brown tissue @ 60m)Vermeij et al., 2002
P-520-570520-570590****Madracis pharensis (green tissue @ 10m)Vermeij et al., 2002
P-520-570520-570590****Madracis pharensis (green tissue @ 20m)Vermeij et al., 2002
P-520-570520-570590-620****Madracis pharensis (green tissue @ 40m)Vermeij et al., 2002
P-520-570520-570590****Madracis pharensis (green tissue @ 60m)Vermeij et al., 2002
P-519-570519-570590-615****Madracis pharensis (green tissue)Vermeij et al., 2002
P-537537590****Madracis pharensis (grey and blue tissue @ 10m)Vermeij et al., 2002
P-532532*****Madracis pharensis (red tissue @ 10m)Vermeij et al., 2002
P-587587609****Madracis pharensis @  40m (brown tissue)Vermeij et al., 2002
P-560560590****Madracis pharensis @ 10m (grey tentacles)Vermeij et al., 2002
P-519-559519-559590****Madracis senariaVermeij et al., 2002
P-520-558520-558590****Madracis senariaVermeij et al., 2002
P-561561587-616****Madracis senaria @ 40mVermeij et al., 2002
P-559559590****Madracis senaria @ 60mVermeij et al., 2002
P-487487515575***Manicina areolataMazel, 1995
P-487487515****Meandrina meandritesMazel,1995
P-480480*****Montastraea annularisManica and Carter, 2000
P-484484499****Montastraea annularisMazel, 1995
P-486486*****Montastraea annularisMazel, 1997
P-503503479578***Montastraea annularisMazel, 1995
P-510510479****Montastraea annularisMazel, 1995
P-515515**505**Montastraea annularisMazel, 1997; Manica & Carter, 2000
P-517517551483***Montastraea annularisMazel, 1995
P-486486**440**Montastraea cavernosaLesser et al., 2000
P-505505*****Montastraea cavernosaKelmanson & Matz, 2003
P-510510**440**Montastraea cavernosaMazel et al., 2003
P-510-520510-520**440**Montastraea cavernosaLesser et al., 2000
P-514514*****Montastraea cavernosaKelmanson & Matz, 2003
P-518518*****Montastraea cavernosaKelmanson & Matz, 2003
P-519519**~505**Montastraea cavernosaKelmanson & Matz, 2003
P-522522*****Montastraea cavernosaKelmanson & Matz, 2003
P-575575~630*~525~570*Montastraea cavernosaMazel, 1997
P-516516**506~480*Montastraea cavernosa (=mc2/3/4 – see P-515)Labas et al., 2002
P-582582630*508572*Montastraea cavernosa (mc1)Kelmanson & Matz, 2003
P-515515±3**505±3**Montastraea cavernosa (mc2/3/4)Kelmanson & Matz, 2003
P-495495**435**Montastraea cavernosa (mc5)Kelmanson & Matz, 2003
P-507507**495**Montastraea cavernosa (mc6)Kelmanson and Matz, 2003
P-580580520*508572*Montastraea cavernosa (mcavRFP)Labas et al., 2002
P-510-623510-623*****Montastraea cavernosa @ 40mVermeij et al., 2002
P-534534593****Montastraea cavernosa @ 60mVermeij et al., 2002
P-486486**440**Montastraea faveolataLesser et al., 2000
P-510-520510-520**440**Montastraea faveolataLesser et al., 2000
P-575575**506555*Montipora (digitata/angulata)Mazel, unpublished
P-485485**420460465Montipora caliculataDove et al., 2001
P-490490**420-450**Montipora monasteriataDove et al., 2001
P-610610**570**Montipora monasteriataDove et al., 2001
P-620620**440**Montipora sp.Kogure et al., 2006
P-515515*****Mycetophyllia lamarckianaMazel, 1997
P-515515**~494**Mycetophyllia sp.Fux and Mazel, unpublished
P-575575**~569~540~500Phycoerythrin within symbiotic cyanobacteria in coralMazel et al., 2004
P-462(?)462*****Platygyra daedaleaSalih et al., 2000
P-505505**492**Plesiastrea verisporaDove et al., 2001
P-574574550****Plesiastrea verisporaDove et al., 2001
P-472472*****Plesiastrea verispora (blue morph)Salih et al., 2004
P-473473*****Plesiastrea verispora (blue morph)Salih et al., 2004
P-479479*****Plesiastrea verispora (blue morph)Salih et al., 2004
P-485485*****Plesiastrea verispora (blue morph)Salih et al., 2004
P-488488*****Plesiastrea verispora (blue morph)Salih et al., 2004
P-489489*****Plesiastrea verispora (blue morph)Salih et al., 2004
P-495495*****Plesiastrea verispora (blue morph)Salih et al., 2004
P-497497*****Plesiastrea verispora (blue morph)Salih et al., 2004
P-501501*****Plesiastrea verispora (blue morph)Salih et al., 2004
P-512512*****Plesiastrea verispora (blue morph)Salih et al., 2004
P-540540*****Plesiastrea verispora (blue morph)Salih et al., 2004
P-580580*****Plesiastrea verispora (blue morph)Salih et al., 2004
P-620620*****Plesiastrea verispora (blue morph)Salih et al., 2004
P-477477*****Plesiastrea verispora (blue)Salih et al., 2004
P-482482*****Plesiastrea verispora (blue)Gilmore et al., 2003
P-492492*****Plesiastrea verispora (blue)Salih et al., 2004
P-473473*****Plesiastrea verispora (green morph)Salih et al., 2004
P-479479*****Plesiastrea verispora (green morph)Salih et al., 2004
P-485485*****Plesiastrea verispora (green morph)Salih et al., 2004
P-489489*****Plesiastrea verispora (green morph)Salih et al., 2004
P-495495*****Plesiastrea verispora (green morph)Salih et al., 2004
P-497497*****Plesiastrea verispora (green morph)Salih et al., 2004
P-503503*****Plesiastrea verispora (green morph)Salih et al., 2004
P-508508*****Plesiastrea verispora (green morph)Salih et al., 2004
P-511511*****Plesiastrea verispora (green morph)Salih et al., 2004
P-512512*****Plesiastrea verispora (green morph)Salih et al., 2004
P-515515*****Plesiastrea verispora (green morph)Gilmore et al., 2003
P-515515*****Plesiastrea verispora (green morph)Salih et al., 2004
P-518518*****Plesiastrea verispora (green morph)Salih et al., 2004
P-540540*****Plesiastrea verispora (green morph)Salih et al., 2004
P-580580*****Plesiastrea verispora (green morph)Salih et al., 2004
P-620620*****Plesiastrea verispora (green morph)Salih et al., 2004
P-440440**358**Pocillopora damicornisDove et al., 2001
P-499499**484**Pocillopora damicornisDove et al., 2001
P-508508**440450*Pocillopora damicornisApprill, 2003
P-516516**486**Pocillopora damicornisSalih et al., 2000
P-625625**570**Pocillopora damicornisDove et al., 2001
P-530530**450***Porites astreoidesMazel, 2003
P-620620**490**Porites astreoidesMazel et al., 2003
P-496496*****Porites cylindricaSalih et al., 2000
P-508508*****Porites cylindricaSalih et al., 2000
P-485485**420**Porites murrayensisDove et al., 2001
P-550550**530**Porites murrayensisDove et al., 2001
P-625625**570**Porites murrayensisDove et al., 2001
P-508508**500**Ptilosarcus sp.Labas et al., 2002
P-510510**498**Renilla muelleri (sea pansy)Labas et al., 2002
P-510510*****Ricordea floridaMazel, 1995
P-513513*****Ricordea floridaMazel, 1995
P-517517574*506566*Ricordea floridaLabas et al., 2002
P-517517**506**Ricordea floridaMazel, 1995
P-518518**508475*Ricordea floridaLabas et al., 2002
P-518518*****Ricordea floridaMazel, 1995
P-520520*****Ricordea floridaMazel, 1995
P-573573510****Ricordea floridaMazel, 1995
P-574574517*506566*Ricordea floridaLabas et al., 2002
P-587-590587-590*****Ricordea florida (mouth)Mazel, 1995
P-515515*****Ricordea sp.Mazel, 1995
P-506506**497**Scolymia cubensis 1Labas et al., 2002
P-506506**497**Scolymia cubensis 2Labas et al., 2002
P-483483511****Scolymia sp.Mazel, 1995
P-483483511576***Scolymia sp.Mazel, 1995
P-484484512****Scolymia sp.Mazel, 1995
P-484484512576***Scolymia sp.Mazel, 1995
P-515515*****Scolymia sp.Mazel, 1997
P-575575630*520**Scolymia sp.Mazel et al., 2003
P-482482462****Seriatopora hystrixDove et al., 2001
P-503503535****Siderastrea radiansMazel, 1995
P-503503535****Stoichactis sp.Mazel, 1995
P-582582**558**Trachyphyllia geoffroyiAndo et al., 2002
P-576576*****Zoanthus – Mature form of P-522Ianushevich et al., 2003
P-506506**496~440*Zoanthus 1Matz et al., 1999
P-538538~580*528494*Zoanthus 2Matz et al., 1999
P-506506**494**Zoanthus sp.Yanushevich et al., 2002
P-538538**525494*Zoanthus sp.Yanushevich et al., 2002
*Isolated pigments – See Comments in text.
**A good example of FRET – several pigments in
A. millepora  ‘relay’ excitation to P-593

 

Table 2. A listing of pigments (by emission spectra) of pigments discussed in Part 1 of this series.
PigmentEmission23Excitation23Found in:Reference
P-400400**345**Acropora horridaDove et al., 2001
P-409409*****Acropora nastuaGilmore et al., 2003
P-440440**358**Pocillopora damicornisDove et al., 2001
P-445445**340**Acropora asperaDove et al., 2001
P-446(?)446**380**Leptoseris fragilisSchlichter et al., 1985
P-462(?)462*****Platygyra daedaleaSalih et al., 2000
P-472472*****Plesiastrea verispora (blue morph)Salih et al., 2004
P-473473*****Plesiastrea verispora (green morph)Salih et al., 2004
P-473473*****Plesiastrea verispora (blue morph)Salih et al., 2004
P-476476510575500475*Acropora aspera (orange band I)*Papina et al., 2002
P-477477*****Plesiastrea verispora (blue)Salih et al., 2004
P-478478510575501475*Acropora aspera (orange band II)*Papina et al., 2002
P-479479*****Plesiastrea verispora (blue morph)Salih et al., 2004
P-479479*****Plesiastrea verispora (green morph)Salih et al., 2004
P-480480510*481**Acropora aspera (green band)*Papina et al., 2002
P-480480515*470504*Acropora tenuis (green band)*Papina et al., 2002
P-480480*****Goniopora tenuidensSalih et al., 2004
P-480480*****Montastraea annularisManica and Carter, 2000
P-482482*****Plesiastrea verispora (blue)Gilmore et al., 2003
P-482482515*451430*Acropora nastuaPapina et al., 2002
P-482482462****Seriatopora hystrixDove et al., 2001
P-482482515*452500425Acropora secale (green band)*Papina et al., 2002
P-483483**572454510Lobophyllia hemprichii (red)Salih et al., 2004
P-483483**443**Discosoma striataMatz et al., 1999
P-483483511****Scolymia sp.Mazel, 1995
P-483483**427451*Acropora nastua (green band)*Papina et al., 2002
P-483483511576***Scolymia sp.Mazel, 1995
P-484484**456**Clavularia sp.Matz et al., 1999
P-484484499****Montastraea annularisMazel, 1995
P-484484512****Scolymia sp.Mazel, 1995
P-484484515*450502*Acropora secalePapina et al., 2002
P-484484512576 **Scolymia sp.Mazel, 1995
P-485485**420**Acropora horridaDove et al., 2001
P-485485*****Plesiastrea verispora (blue morph)Salih et al., 2004
P-485485*****Plesiastrea verispora (green morph)Salih et al., 2004
P-485485**420460465Montipora caliculataDove et al., 2001
P-485485517555465505*Acropora tenuisPapina et al., 2002
P-485485**420**Porites murrayensisDove et al., 2001
P-486486**384**Acropora nobilisSalih et al., 2000
P-486486**458**Anemonia majanoMatz et al., 1999
P-486486**426**Agaricia sp.Mazel, 1997
P-486486**~448**Diploria labyrinthiformisFux and Mazel, unpublished
P-486486*****Montastraea annularisMazel, 1997
P-486486**440**Montastraea faveolataLesser et al., 2000
P-486486**440**Montastraea cavernosaLesser et al., 2000
P-487487517****Acropora cervicornisMazel,1995
P-487487517****Eusmilia fastigataMazel, 1995
P-487487515575***Manicina areolataMazel, 1995
P-487487515****Meandrina meandritesMazel,1995
P-488488*****Plesiastrea verispora (blue morph)Salih et al., 2004
P-489489*****Plesiastrea verispora (blue morph)Salih et al., 2004
P-489489*****Plesiastrea verispora (green morph)Salih et al., 2004
P-490490**425**Acropora digitiferaDove etal., 2001
P-490490**405**Acropora milleporaCox and Salih, 2005
P-490490514*480501*Acropora asperaPapina et al., 2002
P-490490**420-450**Montipora monasteriataDove et al., 2001
P-492492*****Plesiastrea verispora (blue)Salih et al., 2004
P-494494*****Goniopora tenuidensSalih et al., 2004
P-495495**435**Montastraea cavernosa (mc5)Kelmanson & Matz, 2003
P-495495*****Plesiastrea verispora (blue morph)Salih et al., 2004
P-495495*****Plesiastrea verispora (green morph)Salih et al., 2004
P-495495*****Acropora digitiferaGilmore et al., 2003
P-495495**472**Acropora sp.Karasawa et al., 2003
P-496496*****Porites cylindricaSalih et al., 2000
P-496496**399482*Condylactis giganteaLabas et al., 2002
P-497497527****Condylactis giganteaMazel, 1995
P-497497*****Plesiastrea verispora (blue morph)Salih et al., 2004
P-497497*****Plesiastrea verispora (green morph)Salih et al., 2004
P-497497527****Agaricia sp.Mazel, 1995
P-499499**480403278Anemonia sculataWiedenmann et al., 2002
P-499499**484**Pocillopora damicornisDove et al., 2001
P-500500**475**Discosoma sp.Gross et al., 2000
P-500500**405481*Heteractis crispiaLabas et al., 2002
P-500500**480**Acropora asperaDove et al., 2001
P-501501*****Plesiastrea verispora (blue morph)Salih et al., 2004
P-503503479578***Montastraea annularisMazel, 1995
P-503503535****Siderastrea radiansMazel, 1995
P-503503535****Stoichactis sp.Mazel, 1995
P-503503*****Plesiastrea verispora (green morph)Salih et al., 2004
P-504504**405**Acropora milleporaCox and Salih, 2005
P-505505**492**Plesiastrea verisporaDove et al., 2001
P-505505**492**Galaxea fascicularisKarasawa et al., 2003
P-505505*****Montastraea cavernosaKelmanson & Matz, 2003
P-506506**497**Scolymia cubensis 1Labas et al., 2002
P-506506**497**Scolymia cubensis 2Labas et al., 2002
P-506506**494**Zoanthus sp.Yanushevich et al., 2002
P-506506**496~440*Zoanthus 1Matz et al., 1999
P-507507536575***Favia fragumMazel, 1995
P-507507**495**Montastraea cavernosa (mc6)Kelmanson and Matz, 2003
P-508508*****Plesiastrea verispora (green morph)Salih et al., 2004
P-508508**440450*Pocillopora damicornisApprill, 2003
P-508508*****Porites cylindricaSalih et al., 2000
P-508508**494**DendronephthyaLabas et al., 2002
P-508508**500**Ptilosarcus sp.Labas et al., 2002
P-508-620508-620*****Agaricia agaricites @ 60mVermeij et al., 2002
P-509509*****Goniopora tenuidensSalih et al., 2004
P-509509**397**Aequoria victoriaTsien, 1998
P-510510479****Montastraea annularisMazel, 1995
P-510510*****Ricordea floridaMazel, 1995
P-510510**498**Renilla muelleri (sea pansy)Labas et al., 2002
P-510510**490420*Heteractis magnificaTu et al., 2003
P-510510**440**Montastraea cavernosaMazel et al., 2003
P-510-520510-520**440**Montastraea faveolataLesser et al., 2000
P-510-520510-520**440**Montastraea cavernosaLesser et al., 2000
P-510-623510-623*****Montastraea cavernosa @ 40mVermeij et al., 2002

Transcribing the charts from various papers has been
time-consuming work. The charts and the information contained
within these articles are fine for our casual purposes. In all
cases, the original papers should be reviewed for exact graphical
data should requirements go beyond ‘aquarium’
use.

Writing this series of articles has been tedious –
sometimes frustrating. It includes information I’ve
gathered over the last 20 years (see the Reference list at the
end of this article). I have tried to ‘weed through’
as many references as I can, looking for those one or two
pertinent sentences that are often buried in pages of important,
yet irrelevant, material. More often that not, what is of extreme
importance to the genetic engineer is of little practical use to
us as reef hobbyists. These articles are as complete as I can
make them – there is always the temptation to download just
one more paper, send one more email, research one
‘last’ angle. The excitement of discovery is
addictive, yet at some point I must temporarily conclude the
work, knowing the paper will be dated by the time I finish typing
this sentence.

This time, we’ll examine pigments with emissions up to
the true green portion (P-510) of the spectrum. With that said,
let’s get started!

Violet and Blue Pigments

Figure6.jpg

Figure 6. A beautiful Acropora specimen from the
aquarium of LA hobbyist Dave Botwin. Photo by the author.

Relatively little is known about blue fluorescent coral
pigments. The biomedical field has little interest in these since
they do not work well in many studies due to cellular
autofluorescence. In additional, the excitation wavelengths can
harm living cells. Solving the issue of blue fluorescence (or
blue non-fluorescence for that matter) will probably be left in
the hands of coral researchers interested in these
pigments’ possible ecological functions.

It is interesting to note that excitation wavelengths for
P-400, P-440 and P-445 absorb some of the ultraviolet radiation
in a range that natural-occurring UV sunscreens do not- MAAs or
mycosporine-like amino acids absorb UV energy up to ~360nm
(Bandaranayake, 1998).

P-400

Host: Acropora horrida

  • Excitation: 345nm
  • Emission: 400nm
  • Stokes shift: 55nm
  • Reference: Dove et al., 2001
  • Comments: No fluorescent quantum yield is
    available for this pigment. Violet fluorescence is uncommon in anthozoans.
    This pigment’s absorption suggests exposure to UV radiation
    (such as that from an unshielded metal halide lamp) would result
    in maximum fluorescence. Salih et al., 1998 state that certain
    Acropora species including, among others, A.
    horrida
    and A. tortuosa, sometimes are devoid of
    zooxanthellae within their polyps (the symbiotic dinoflagellates
    are concentrated in other tissues). In these cases, the coral
    polyps will not contain fluorescent pigments, but coloration is
    found in areas where zooxanthellae are concentrated. This
    certainly suggests (but does not prove) that the pigments have a
    biological function, perhaps as a natural sunscreen. This
    particular Acropora specimen appeared blue in natural
    light.

P-409

Host: Acropora nastua

  • Excitation: Not listed
  • Emission: 495nm
  • Stokes shift: N/A
  • Reference: Gilmore et al., 2003
  • Comments: This specimen appears brown with blue tips in
    natural light and was collected at a depth of 1.5m at Cape Maeda,
    Okinawa, Japan.

P-440

Host: Pocillopora damicornis

  • Excitation: 358nm
  • Emission: 440nm
  • Stokes shift: 82nm
  • Reference: Dove et al., 2001
  • Comments: This coral specimen appeared pink in natural light.
    Maximum absorption of coral tissue is 560nm, indicating the
    presence of a non-fluorescent chromoprotein (Dove et al., 1995).
    See Figure 7.
Figure7.JPG

Figure 7. Excitation and emission spectra of a blue
fluorescent protein found in the stony coral Pocillopora
damicornis
. After Dove et al., 2001.

P-445

Host: Acropora aspera

  • Excitation: 340nm
  • Emission: 445nm
  • Stokes shift: 105nm
  • Reference: Dove et al., 2001
  • Comments: Maximum absorption of coral tissue, zooxanthellae
    and pigments is at 580nm (data not shown) due to the presence of
    a non-fluorescent chromoprotein – pocilloporan. This Acropora specimen
    was blue in color. See Figure 8.
Figure8.JPG

Figure 8. Fluorescent blue excitation and emission spectra
from Acropora aspera. After Dove et al.,
2001.

Figure9.JPG

Figure 9. An apparent example of fluorescent coupling where
emission spectra overlap the excitation spectra of the
‘next’ pigment. Although this coral appears blue to
the eye in natural light (due to the presence of a
non-fluorescent protein), its peak fluorescent emission is in the
blue-green portion of the spectrum.

P-446 (?)

Although Schlichter et al. (1985, 1986) do not specifically
mention the maximum fluorescence wavelength of their deep-water
Leptoseris fragilis, it appears as if the fluorescent peak
is close to 446nm (when variously excited by ultraviolet
radiation at 380 and 390nm, violet at 400nm and blue at 430nm).
The shoulder at ~380nm in Figure 10 is caused by the excitation
light.

Figure10.JPG

Figure 10. Emission spectrum of a deep-water coral
Leptoseris fragilis. The theory of a fluorescent pigment
enhancing photosynthesis was developed around this data. The peak
in the emission at 380nm is actually due to the UV excitation
source (this portion of the smoothed line has been color-coded
violet for easy differentiation. After Schlichter et al.
1986).

P-462 (?)

Salih et al., 2000 note a ‘dense blue’ layer of
fluorescent pigment in the oral disk of a shallow-water stony
coral Platygyra daedalea. No exact emission wavelength is
listed, although it appears to be ~462nm in one of the
article’s charts.

P-472

Host: Plesiastrea verispora

  • Excitation: 330-380nm
  • Emission: 472nm
  • Stokes shift: Unknown
  • Reference: Salih et al., 2004
  • Comments: This P. verispora appeared blue in natural
    light, and was collected at a depth of 4-8m in Port Jackson,
    Australia.

P-473

Host: Plesiastrea verispora

  • Excitation: 330-380nm
  • Emission: 473nm
  • Stokes shift: Unknown
  • Reference: Salih et al., 2004
  • Comments: These P. verispora appeared green or blue in
    natural light, and were collected at a depth of 4-8m in Port
    Jackson, Australia.

P-477

Host: Plesiastrea verispora

  • Excitation: 330-380nm
  • Emission: 476nm
  • Stokes shift: Unknown
  • Reference: Salih et al., 2004
  • Comments: This P. verispora appeared blue in natural
    light, and was collected at a depth of 4-8m in Port Jackson,
    Australia.

P-479

Host: Plesiastrea verispora

  • Excitation: 330-380nm
  • Emission: 477nm
  • Stokes shift: Unknown
  • Reference: Salih et al., 2004
  • Comments: These P. verispora appeared green or blue in
    natural light, and were collected at a depth of 4-8m in Port
    Jackson, Australia.

Green-Blue Pigments

P-480

Host: Acropora aspera

  • Excitation: 504nm, with a shoulder at 510nm
  • Emission: 480nm
  • Stokes shift: 70-76nm
  • Reference: Papina et al., 2002
  • Comments: This emission was identified in a ‘green
    band’ from a SDS-PAGE separation.

P-482

Host: Acropora nastua

  • Excitation: 451nm, shoulder at 430nm
  • Emission: 482nm, with a shoulder at 515nm
  • Stokes shift: 31nm
  • Reference: Papina et al., 2002
  • Comments: This Acropora nastua specimen was collected
    off the coast of Okinawa, Japan at a depth of 1.5m in August,
    2001 (Papina et al., 2002). In daylight, the coral appeared brown
    with blue tips (the blue coloration probably due to a
    non-fluorescent chromoprotein). When viewed under a black light,
    the coral fluoresced blue. pH values ranging from 5.0 to 8.0 did
    not affect fluorescent quantum yield or spectral characteristics.
    See Figure 11.
Figure11.JPG

Figure 11. Excitation and emission spectra of a blue
fluorescent pigment. In natural light, this animal appeared brown
with blue tips. After Papina et al., 2002.

Host: Seriatopora hystrix

  • Excitation: 462nm
  • Emission: 482nm
  • Stokes shift: 20nm
  • Reference: Dove et al., 2001
  • Comments: Maximum absorption of coral tissue, zooxanthellae
    and pigments combined is at 560nm (data not shown). This coral
    appeared pink in natural light, due to a non-fluorescent pink
    pigment (one of a group of pigments collectively called
    pocilloporans, Dove et al., 1995). See Figure 12.
Figure12.JPG

Figure 12. This pink Seriatopora hystrix contains a
green-blue fluorescent pigment. After Dove et al., 2001.

Host: Plesiastrea verispora

  • Excitation: Not listed
  • Emission: 482nm
  • Stokes shift: N/A
  • Reference: Gilmore et al., 2003.
  • Comments: ThesePlesiastrea specimens (appearing blue in
    natural light) were collected in 5-9m of water at Port Jackson,
    Australia.

P-483

Host: Scolymia sp.

  • Excitation: Not listed
  • Emission: 483nm
  • Stokes shift: N/A
  • Reference: Mazel, 1995
  • Comments: Emission shoulders at 511nm and 576nm. See Figure
    13.
Figure13.JPG

Figure 13. A fluorescent protein from the stony coral
Scolymia. After Mazel, 1995.

Host: Discosoma striata

  • Excitation: ~458nm
  • Emission: 483nm
  • Stokes shift: 25nm
  • Reference: Matz et al., 1999
  • Comments: This pigment has a quantum yield of 0.46. Brightness
    (relative to the GFP in A. victoria) is 0.50. P-483 is
    structurally similar to P-583 (an orange-red pigment also found
    in some Discosoma specimens. Substitutions of only two
    amino acid residues in the DNA account for the shift of color
    from green-blue to orange-red). This pigment was concentrated in
    blue-green stripes on the animal’s oral disk. It apparently
    was in combination with an unidentified cyan fluorescent
    pigment.

Host: Acropora nastua

  • Excitation: 427nm and 451nm
  • Emission: 483nm
  • Stokes shift: Unknown
  • Reference: Papina et al., 2002
  • Comments: This ‘green band’ was identified as an
    emission in a SDS-PAGE separation of
    pigments.

P-484

Host: Scolymia sp.

  • Excitation: Not listed
  • Emission: 484nm
  • Stokes shift: N/A
  • Reference: Mazel, 1995
  • Comments: Shoulder emissions at 512nm and 576nm. See Figure
    14.

Host: Acropora secale

  • Excitation: 450nm, with an excitation shoulder at 502nm.
  • Emission: 484nm, with a shoulder at 515nm.
  • Stokes shift: 34nm
  • Reference: Papina et al., 2002
  • Comments: Collected in shallow water (1.5m depth) off the
    coast of Okinawa, Japan in August 2001. pH values ranging from
    5.0 to 8.0 did not significantly affect fluorescent quantum yield
    or spectral absorption/emission qualities. Judging from the
    excitation and emission wavelengths, it appears from this pigment
    might be activated (probably by UV or violet/blue light) to
    P-515, which would change the apparent fluorescence from
    green-blue to more of a green coloration. The appearance of this
    coral was brown (due to zooxanthellae) with pink tips (probably
    due to the presence of the chromoprotein CP-560, a
    ‘pocilloporan’).

Host: Montastrea annularis

  • Excitation: Not listed
  • Emission: 484nm, with emission shoulder at 499nm.
  • Stokes shift: N/A
  • Reference: Mazel, 1995
  • Comments:

Host: Clavularia sp

  • Excitation: ~456nm
  • Emission: 484nm
  • Stokes shift: ~28nm
  • Reference: Matz et al., 1999
  • Comments: Fluorescent quantum yield is 0.48. Brightness
    (relative to the emission of A. victoria) is 0.77. As most
    hobbyists know, the intense green fluorescence is concentrated in
    the oral disk and tentacles.
Figure14.JPG

Figure 14. A green-blue pigment from a Caribbean stony coral.
After Mazel, 1995.

Figure15.JPG

Figure 15. Though the emission peak in this Acropora
peaks at the same wavelength as seen in Figure 14, but they do
not seem to be the same pigment. After Papina et al.,
2002.

P-485

Host: Acropora horrida

  • Excitation: 420nm
  • Emission: 485nm
  • Stokes shift: 65nm
  • Reference: Dove et al., 2001; Salih et al., 1999.
  • Comments: Maximum absorption at 579nm (data not shown). Acropora horrida’s polyps
    often do not contain zooxanthellae – they, along with fluorescent coloration,
    are found elsewhere within the coral host’s tissues (Salih
    et al., 1999). This coral is blue in natural light.

Host: Montipora caliculata

  • Excitation: 420nm, 450nm and 465nm
  • Emission: 485nm
  • Stokes shift: 20-65nm
  • Reference: Dove et al., 2001
  • Comments: Maximum absorption at 579nm (data not shown). This
    specimen was colored purple in natural light – probably as
    a result of the presence of a non-fluorescent
    chromoprotein.

Host: Porites murrayensis

  • Excitation: 420nm
  • Emission: 485nm
  • Stokes shift: 65nm
  • Reference: Dove et al., 2001
  • Comments: This coral is purple in color in natural light, with
    a tissue maximum absorption at 576nm (data not shown). The purple
    coloration is likely due to the presence of a non-fluorescent
    chromoprotein.

Host: Plesiastrea verispora

  • Excitation: 330-380nm
  • Emission: 485nm
  • Stokes shift: Unknown
  • Reference: Salih et al., 2004
  • Comments: These P. verispora appeared green or blue in
    natural light, and were collected at a depth of 4-8m in Port
    Jackson, Australia.

P-486

Host: Diploria labrynthiformis

  • Excitation: Not listed
  • Emission: 486nm
  • Stokes shift: N/A
  • Reference: Mazel, unpublished
  • Comments: Specimen collected in the Caribbean.

Host: Agaricia sp.

  • Excitation: 426nm
  • Emission: 486nm
  • Stokes shift: 60nm
  • Reference: Mazel, 1997
  • Comments: See Figure 16.
Figure16.JPG

Figure 16. The fluorescent emission of a Caribbean coral.
After Mazel et al. 1997.

Host: Anemonia majano

  • Excitation: 458nm
  • Emission: 486nm
  • Stokes shift: 28nm
  • References: Matz et al., 1999; Henderson and Remington,
    2005.
  • Comments: Fluorescent quantum yield is 0.24. Relative
    Brightness (GFP from Aequorea victoria as the
    standard) is 0.43. Extracted pigment appears yellow-green in room
    lighting. Light absorbance of the wild type protein is not
    particularly pH-sensitive. Matz et al. mention the bright green
    fluorescent pigment is concentrated in the anemone’s
    tentacle tips. See Figure 17.
Figure17.JPG

Figure 17. A fluorescence most hobbyists would rather not see
– that of the ‘little bastard’ anemone,
Anemonia majano. After Matz et al., 1999.

Host: Acropora nobilis

  • Excitation: 384nm
  • Emission: 486nm
  • Stokes shift: 102nm
  • Reference: Salih et al., 2000
  • Comments: This Acropora specimen appeared blue when
    excited with UV radiation.

See Figure 18.

Figure18.JPG

Figure 18. Fluorescent excitation and emission of the
Indo-Pacific coral Acropora nobilis. After Salih et al.,
2000.

P-487

Host: Acropora cervicornis

  • Excitation: Not listed
  • Emission: 487nm
  • Stokes shift: N/A
  • Reference: Mazel, 1997
  • Comments: Has emission shoulder at 517nm. See Figure
    19.

Host: Eusmilia fastigiata

  • Excitation: Not listed
  • Emission: 487nm
  • Stokes shift: N/A
  • Reference: Mazel, 1997
  • Comments: See Figure 20.

Host: Meandrina meandrites

  • Excitation: Not listed
  • Emission: 487nm
  • Stokes shift: N/A
  • Reference: Mazel, 1997
  • Comments:

Host: Manicina areolata

  • Excitation: Not listed
  • Emission: 487nm
  • Stokes shift: N/A
  • Reference: Mazel, 1997
  • Comments: Emission shoulders were noted at 515nm and 575 nm.
    Though this animal contains various fluorescent compounds, it is
    commonly called the Rose coral for its pink/red
    coloration.
Figure19.JPG

Figure 19. The green-blue emission of the Atlantic coral
Acropora cervicornis. After Mazel et al., 1997.

Figure20.JPG

Figure 20. Emission spectrum of P-487 from a stony coral.
After Mazel, 1997.

P-488

Host: Plesiastrea verispora

  • Excitation: 330-380nm
  • Emission: 488nm
  • Stokes shift: Unknown
  • Reference: Salih et al., 2004
  • Comments: This P. verispora appeared blue in natural
    light, and was collected at a depth of 4-8m in Port Jackson,
    Australia.

P-489

Host: Plesiastrea verispora

  • Excitation: 330-380nm (blue); 450-490nm (green)
  • Emission: 489nm
  • Stokes shift: Unknown
  • Reference: Salih et al., 2004
  • Comments: These P. verispora appeared blue or green in
    natural light, and were collected at a depth of 4-8m in Port
    Jackson, Australia.

P-490

Host: Acropora aspera

  • Excitation: 420-450nm
  • Emission: 490nm
  • Stokes shift: 40-70nm
  • Reference: Dove et al., 2001
  • Comments: This particular coral was blue when viewed in
    natural light. Maximum light absorbance by tissue is ~580nm (data
    not shown). See Figure 21 for excitation and emission data.

Host: Acropora digitifera

  • Excitation: 425nm
  • Emission: 490nm
  • Stokes shift: 70nm
  • Reference: Dove et al., 2001
  • Comments: Coral appeared purple-blue in natural light. Maximum
    absorbance of the tissue is at 578nm (data not shown).

Host: Acropora millepora

  • Excitation: 405nm
  • Emission: 490nm
  • Stokes shift: 85nm
  • Reference: Cox and Salih, 2005.
  • Comments: Excitation via pulsed 405nm laser.

Host: Montipora monasteriata

  • Excitation: 420-450nm
  • Emission: 490nm
  • Stokes shift: 40-70nm
  • Reference: Dove et al., 2001
  • Comments: Coral appeared purple in natural light. Maximum
    tissue absorbance equals 574-578nm (data not
    shown).
Figure21.JPG

Figure 21. One of the more studied corals (pigment-wise)
Acropora aspera. After Dove et al., 2001.

Blue-Green Pigments

Figure22.JPG

Figure 22. Fluorescence of a Hawaiian stony coral
Pavona. Photo by the author.

P-492

Host: Plesiastrea verispora

  • Excitation: 330-380nm
  • Emission: 488nm
  • Stokes shift: Unknown
  • Reference: Salih et al., 2004
  • Comments: This P. verispora appeared blue in natural
    light, and was collected at a depth of 4-8m in Port Jackson,
    Australia.

P-495

Host: Montastraea cavernosa

  • Excitation: 435nm
  • Emission: 495nm
  • Stokes shift: 60nm
  • Reference: Kelmanson and Matz, 2003
  • Comments: M. cavernosa can contain many pigments
    – this one is referred to as mc5. See Figure
    23.
Figure23.JPG

Figure 23. Data for a blue-green pigment found in the stony
coral Montastraea cavernosa. After Kelmanson and Matz,
2003.

Host: Acropora sp.

  • Excitation: 472nm
  • Emission: 495nm
  • Stokes shift: 23nm
  • Reference: Karasawa et al., 2004.
  • Comments: This pigment was found in an unidentified Acropora species
    in the water off Okinawa, Japan. The pigment is concentrated in the polyps.
    It is commercially available as MiCy (Midori-ishi is Japanese for Acropora; Cy is
    for the color –
    cyan).

Host: Acropora digitifera

  • Excitation: Not listed
  • Emission: 495nm
  • Stokes shift: N/A
  • Reference: Gilmore et al., 2003
  • Comments: This specimen appears brown with blue tips in
    natural light and was collected at a depth of 1.5m at Cape Maeda,
    Okinawa, Japan.

Host: Plesiastrea verispora

  • Excitation: 330-380nm (blue); 450-490nm (green)
  • Emission: 489nm
  • Stokes shift: Unknown
  • Reference: Salih et al., 2004
  • Comments: These P. verispora appeared blue or green in
    natural light, and were collected at a depth of 4-8m in Port
    Jackson, Australia.

P-496

Host: Condylactis gigantea

  • Excitation: 399nm, with a shoulder at 482nm.
  • Emission: 496nm
  • Reference: Labas et al., 2002
  • Comments: P-496 is found in the anemone Condylactis
    gigantea
    (Labas 2002). These anemones are popular in aquaria,
    and for good reason. They are easy to maintain in captivity and
    coloration can be spectacular. It is not uncommon to see them
    with green/purple tentacles and contrasting magenta or purple
    tips (Sprung and Delbeek, 1997). The column or stalk can be
    orange or yellow and is due to non-fluorescent chromoproteins.
    However, fluorescence is blue-green. Labas et al. (2002) believe
    the double peaked excitation spectrum (similar to wild-type GFP)
    suggests that the pigment is photoconvertible, though this
    opinion is the subject of debate (Mazel, personal communication;
    Tsien, 1998). See Figure
    24.
Figure24.JPG

Figure 24. P-496 is interesting in that we see evidence
suggesting photoconversion is possible – others disagree.
More research is needed – hobbyists can help! After Labas
et al., 2002.

P-497

Host: Condylactis gigantea

  • Excitation: Not listed
  • Emission: 497nm
  • Stokes shift: N/A
  • Reference: Mazel, 1995
  • Comments: The Atlantic anemone’s fluorescent emission
    also has a shoulder emission at 527nm. See Figure 25.

Host: Agaricia sp.

  • Excitation: Not listed
  • Emission: 497nm
  • Stokes shift: N/A
  • Reference: Mazel, 1995
  • Comments:
Figure25.JPG

Figure 25. Fluorescent emission of the anemone
Condylactis. After Mazel, 1995.

Host: Plesiastrea verispora

  • Excitation: 330-380nm (blue); 450-490nm (green)
  • Emission: 489nm
  • Stokes shift: Unknown
  • Reference: Salih et al., 2004
  • Comments: These P. verispora appeared blue or green in
    natural light, and were collected at a depth of 4-8m in Port
    Jackson, Australia.

P-499: Precursor to P-522

Host: Anemonia sculata

  • Excitation: 278, 403 and 480nm
  • Emission: 499nm, with shoulder at 522nm.
  • Stokes shift: 19nm
  • Reference: Wiedenmann et al., 2000; Labas et al., 2002;
    Verkhusha et al., 2001
  • Comments: The green fluorescent protein is apparently the
    immature form of P-522.

Host: Pocillopora damicornis

  • Excitation: 484nm
  • Emission: 499nm
  • Stokes shift: 55nm
  • Reference: Dove et al., 2001
  • Comments: This P. damicornis appeared pink in natural
    light. Maximum tissue absorption equals 560nm (data not shown),
    but see Figure 26 for fluorescence information.
Figure26.JPG

Figure 26. Information for a blue-green pigment found in
Pocillopora. In natural light the coral appears pink.
After Dove et al., 2001.

P-500: Possible Precursor of Pigment 580

Host: Acropora aspera

  • Excitation: 480nm
  • Emission: 500nm
  • Stokes shift: 20nm
  • Reference: Dove et al., 2001
  • Comments: Maximum tissue absorbance is at 580nm (data not
    shown). This coral appears blue in natural light.

Host: Heteractis crispa

  • Excitation: 405nm, with a shoulder at 481nm.
  • Emission: 500nm
  • Stokes shift: 95nm
  • Reference: Labas et al., 2002
  • Comments: Double peaked excitation suggests that
    photoconversion is possible (Labas et al., 2002) – again,
    this is a matter of debate in the scientific community. Heteractis also
    has a non-fluorescent chromoprotein.

Host: Discosoma sp.

  • Excitation: 475nm
  • Emission: 500nm
  • Stokes shift: 25nm
  • Reference: Gross et al., 2000.
  • Comments: Minor substitutions in the residue via mutagenesis
    prevent maturation.
Figure27.JPG

Figure 27. A second example of excitation and emission spectra
suggesting that photoconversion is possible. This pigment is
found in the anemone Heteractis. After Labas et al., 2002.
More work is needed to confirm possibly color shifts – the
opinion of Labas et al. is not universally accepted.

P-501

Host: Plesiastrea verispora

  • Excitation: 450-490nm
  • Emission: 501nm
  • Stokes shift: Unknown
  • Reference: Salih et al., 2004
  • Comments: This Plesiastrea appeared blue (likely due to
    the presence of a pocilloporan protein) and was collected in 4-8m
    of water at Port Jackson, Australia.

P-503

Host: Montastrea annularis

  • Excitation: Not listed
  • Emission: 503nm, with shoulders at 479nm and 578nm
  • Stokes shift: N/A
  • Reference: Mazel, 1995
  • Comments:

Host: Siderastrea radians

  • Excitation: Not listed
  • Emission: 503nm
  • Stokes shift: N/A
  • Reference: Mazel, 1995
  • Comments:

Host: Stoichactis sp.

  • Excitation: Not listed
  • Emission: 503nm, shoulder at 535nm
  • Stokes shift: N/A
  • Reference: Mazel, 1995
  • Comments: Giant Carpet anemones (Stoichactis) are
    green, purple, pink, blue or fluorescent reddish pink- Sprung and
    Delbeek, 1997.

Host: Plesiastrea verispora

  • Excitation: 450-490nm
  • Emission: 503nm
  • Stokes shift: Unknown
  • Reference: Salih et al., 2004
  • Comments: This Plesiastrea appeared green in natural
    light and was collected in 4-8m of water at Port Jackson,
    Australia.

P-504

Host: Acropora millepora

  • Excitation: 405nm
  • Emission: 504nm
  • Stokes shift: 99nm
  • Reference: Cox and Salih, 2005.
  • Comments: Excitation via pulsed 405nm laser.

P-505

Host: Montastraea cavernosa

  • Excitation: Not listed
  • Emission: 505nm
  • Stokes shift: N/A
  • Reference: Kelmanson and Matz, 2003.

Host: Plesiastrea verispora

  • Excitation: 492nm
  • Emission: 505nm
  • Stokes shift: 13nm
  • Reference: Dove et al., 2001
  • Comments: This Plesiastrea specimen appeared green in
    natural light. See Figure 28.
Figure28.JPG

Figure 28. Although green in color under natural light, this
stony coral will visibly fluoresce blue-green under the proper
conditions. After Dove et al., 2001.

Host: Galaxea fascicularis

  • Excitation: 492nm
  • Emission: 505nm
  • Stokes shift: 13nm
  • Reference: Karasawa et al., 2003.
  • Comments: This pigment is commercially available as
    “Azami-Green” (the Japanese name for Galaxea is
    “Azami-Sango”). Galaxea specimens, despite their
    aggressive nature through use of long sweeper tentacles, are
    often kept in reef aquaria. See Figure 29.
Figure29.JPG

Figure 29. Blue-green fluorescence of the ‘nasty’
stony coral Galaxea. After Karasawa et al., 2003.

P-506

Host: Zoanthus 1

  • Excitation: 492nm
  • Emission: 506, with shoulder at ~540nm
  • Stokes shift: 14nm
  • Reference: Matz et al., 1999.
  • Comments: Fluorescent quantum yield = 0.63; Relative
    Brightness =1.02 (meaning it is just slightly brighter than the A. victoria GFP
    reference standard). See Figure 30.

Host: Scolymia cubensis

  • Excitation: 497nm
  • Emission: 506nm
  • Stokes shift: 9nm
  • Reference: Labas et al., 2002.
  • Comments: Listed as GFPs 1 and 2 from Scolymia cubensis by Labas
    et al., 2002.
Figure30.JPG

Figure 30. One of several similar fluorescent proteins from
Zoanthus. After Matz et al., 1999.

P-507

Host: Favia fragum

  • Excitation: Not listed
  • Emission: 507nm, shoulders at 536nm and 575nm
  • Stokes shift: N/A
  • Reference: Mazel, 1995
  • Comments:

Host: Montastraea cavernosa

  • Excitation: Not listed
  • Emission: 507nm
  • Stokes shift: N/A
  • Reference: Kelmanson and Matz, 2003.
  • Comments: M. cavernosa can contain many pigments
    – this one is referred to as mc6. See Figure
    31.
Figure31.JPG

Figure 31. One of the many fluorescence colors of the stony
coral Montastraea. After Kelmanson and Matz, 2003.

P-508: Precursor of Pigment 575

Host: Dendronephthya sp.

  • Excitation: 494nm
  • Emission: 508nm
  • Stokes shift: 14nm
  • Reference: Labas et al., 2002
  • Comments: P-508 matures from green to red under intense blue
    light (Labas 2002) even in anaerobic conditions, but not in the
    dark – light mediates the change in the process known as
    photoconversion (Nienhaus et al., 2005). It is interesting that a
    fluorescent pigment found in a coral void of zooxanthellae
    (especially one thought capable of photoconversion when exposed
    to blue light) is considered by some to act as a photoprotectant.
    This pigment is commercially available as Dendra
    (‘Dend’ for Dendronephthya and
    ‘ra’ for red activatable). See Figure 32.

Host: Porites cylindrica

  • Excitation: Not listed
  • Emission: 508nm
  • Stokes shift: N/A
  • Reference: Salih et al., 2000
  • Comments: Emission shoulders at 496 and 508nm (Salih et al.,
    2000).

Host: Ptilosarcus guernyi (Sea Pen)

  • Excitation: 500nm
  • Emission: 508nm
  • Stokes shift: 8nm
  • Reference: Labas et al., 2002; Peele et al., 2001
  • Comments: These Sea Pens are often found on soft or muddy
    bottoms at depths to ~75m. This is yet another example of a
    fluorescent pigment within an azooxanthellate cnidarian. This GFP
    is believed by some to be of very early origin, and perhaps a
    precursor to many of the fluorescent pigments seen
    today.
Figure32.JPG

Figure 32. The soft coral Dendronephthya is
non-photosynthetic but contains a protein claimed by some to act
as a photoprotectant. After Labas et al., 2002.

P-509: The ‘Original’ Green Fluorescent
Protein

The ‘original’ green fluorescent protein (the
widely accepted name and abbreviated as GFP) was isolated from
tissues of the jellyfish Aequorea victoria and, later, the
Sea Pen Renilla reniformis. Although it was described
early on, P-509 has not been placed with sufficient confidence
within the coral pigment phylogenetic tree and is therefore not
positively described as an ancestreal form (Labas et al., 2002).
Years of attempts to engineer P-509 to fluoresce far-red shifted
wavelengths have failed. However, P-509’s absorption
characteristics are altered by exposure to wavelengths 400nm and
shorter (meaning ultraviolet radiation). With this pigment, we
have more information on the potential effects of light energy at
relatively narrow bandwidth on the spectral characteristics.
Absorbance increases at 478nm (and decreases at 398nm) when
irradiated with monochromatic light of 398nm. Interestingly,
absorbance increases at 478nm when the pigment is excited by the
same wavelength. This process can take hours, and recovery is at
about 60% after 24 hours (Chattoraj et al., 1996; See Figure
34.).

Host: Aequorea victoria (Jellyfish)

  • Excitation: 397nm
  • Emission: 509nm
  • Stokes shift: 112nm
  • Reference: Tsien, 1998.
  • Comments: Discovered by Osamu Shimomura et al. in the
    jellyfish Aequorea victoria during the early 1960’s
    (Shimomura et al., 1962). Formation of the fluorophore requires
    molecular oxygen and apparently does not require enzymes or
    cofactors (Heim et al., 1994). Emission maximum is sometimes
    listed as 508nm. Emission is 503nm when excited by green-blue
    light at 476nm. See Figure 33.

Host: Renilla reniformis (Sea Pansy)

  • Excitation: 498nm
  • Emission: 509nm
  • Stokes shift: 11nm
  • Reference: Labas et al., 2002
  • Comments: Emission variously listed as 509nm and
    510nm.
Figure33.JPG

Figure 33. P-509, one of the first identified fluorescent
pigments. From Tsien, 1998.

Figure34.JPG

Figure 34. Ultraviolet irradiation (at 398nm) causes a shift
in absorbance of Aequorea victoria’s GFP. See text
for details. After Chattoraj et al., 1996.

P-510

Host: Montastraea annularis

  • Excitation: 440nm
  • Emission: 510nm
  • Stokes shift: 70nm
  • Reference: Mazel et al., 2003.
  • Comments: Shoulder at 479 nm. See Figure 35.

Host: Ricordea florida

  • Excitation: Not listed
  • Emission: 510nm
  • Stokes shift: N/A
  • Reference: Mazel, 1995
  • Comments: Shoulder at 479 nm.
Figure35.JPG

Figure
35. Montastraea cavernosa fluoresces blue-green. After Mazel et al.,
2003.

The quick glance at Pigment 510 concludes Part 1 of this
series. Next time, we’ll examine some of the truly green
and other variously colored fluorescent pigments. We’ll
also see some confirmed examples of photoconversion. If
you’re curious about coloration, this promises to get
pretty interesting.

For convenience (mostly mine), I have chosen to list below all
references cited in this series of articles. If you’re
interested in discussing coral coloration, please email me at
RiddleLabs@aol.com.

Acknowledgement

Many mahalos to Charles Mazel for his time and helpful
comments during the preparation of this article. See his website
at www.nightsea.com.

References and Further Reading

  1. Ando, R., H. Hama, M. Yamamoto-Hino, H. Mizuno, and A.
    Miyawaki, 2002. An optical marker based on the UV-induced
    green-to-red photoconversion of a fluorescent protein. Proc.
    Natl. Acad. Sci. USA, 99(20):12651-12656.
  2. Ando, R., H. Mizuno and A. Miyawaki, 2004. Regulated fast
    nucleocytoplasmic shuttling observed by reversible protein
    highlighting. Science, 306:1370-1373.
  3. Andresen, M., M. Wahl, A. Stiel, F. Gräter, L.
    Schäfer, S. Trowitzsch, G. Weber, C. Eggeling, H.
    Grubmüller, S. Hell and S. Jakobs, 205. Structure and
    mechanism of the reversible photoswitch of a fluorescent
    protein. Proc. Natl. Acad. Sci. USA, 102, 37:13070-13074.
  4. Apprill, A., 2003. Spectral characteristics and genetic
    expression of green fluorescent proteins in Hawaiian corals.
    In: Molecular Biology of Corals: Results of 2002 Edwin W.
    Pauley Summer Program in Marine Biology
    , E. Cox and T.
    Lewis, eds. University of Hawaii HIMB Technical Report No.
    43:6-13.
  5. Baird, G., D. Zacharias and R. Tsien, 2000. Biochemistry,
    mutagenesis, and oligomerization of DsRed, a red fluorescent
    protein from coral. Proc. Natl. Acad. Sci. USA,
    97(22):11984-11989.
  6. Bandaranayake, W., 1998. Mycosporines: Are they
    nature’s sunscreens? National Product Review, 1998.
    159-172.
  7. Bingman, C., 1995. Green-fluorescent protein: a model for
    coral host fluorescent proteins? Aquarium Frontiers, 2(3):
    6-9.
  8. Bingman, C., 1999. Biochemistry of Aquaria: Coral
    Fluorescence – An Update.

    http://www.reefs.org/library/aquarium_frontiers/index.html
  9. Blundell, A., 2005. Lateral Lines: The Seen and Unseen
    World of Coral Fluorescence.
    http://www.advancedaquarist.com/2005/2/lines/
  10. Bulina, M., D. Chudakov, N. Mudrik, and K. Lukyanov, 2002.
    Interconversion of Anthozoa GFP-like fluorescent and
    non-fluorescent proteins by mutagenesis. BMC Biochem., 24;
    3(1):7.
  11. Bulina, M., K. Lukyanov, I. Yampolsky, D. Chudakov, D.
    Staroverov, A. Shcheglov, N. Gurskaya and S. Lukyanov, 2004.
    New class of blue animal pigments based on frizzled and kringle
    protein domains. J. Biol. Chem., 279(42):43367-43370.
  12. Burr, A., P. Hunt, D. Wagar, S. Dewilde, M. Blaxter, J.
    Vanfleteren and L. Moens, 2000. A hemoglobin with an optical
    function. J. Biol. Chem., 275(7): 4810-4815.
  13. Calfo, A., 2005. Magnificent fluorescence! Aquaristic
    perspectives.
    http://reefkeeping.com/issues/2005-11/ac/index.php
  14. Chattoraj, M., B. King, G. Bublitz and S. Boxer, 1996.
    Ultra-fast excited state dynamics in green fluorescent protein:
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Categories:
  Advanced Aquarist, Advanced Aquarist
Dana Riddle
About

 Dana Riddle

  (120 articles)

I have been an aquarist since 1964 and a reef hobbyist since the mid-1980’s. I am the owner of a small laboratory (Riddle Aquatic Laboratories) that specializes in investigation of interactions between light and water motion & photosynthetic organisms (especially corals). The results of this research, resulting in almost 250 articles, have been published in Advanced Aquarist Online, Aquarium Frontiers, Koralle, Freshwater and Marine Aquarium, The Breeders’ Registry, Aquarium Fish, Marine Fish Monthly and others. My first article was published in a 1984 SeaScope and relayed my experiences with a refugium – an idea that would catch fire about a decade later. I have had the honor of making over 60 presentations to various groups, including national conferences such as the Marine Aquarium Conference of North America (MACNA) International Marine Aquarium Conference (IMAC), PetsFestival (Italy), regional conferences, and local clubs. I received the Marine Aquarium Society of North America (MASNA) Aquarist of the Year Award in 2011 at the MACNA conference in Des Moines.

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