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Marine Research Report

Bioluminescence – a source of marine energy?

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Bioluminescence – a source of

marine energy?

Dr. Jessica Craig

Prof. Imants G. Priede

February 2012

Oceanlab®, University of Aberdeen,

Main Street, Newburgh, Aberdeenshire,

U.K., AB41 6AA

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© Crown Copyright 2012

978-1-906410-32-2

Published by The Crown Estate on behalf of the Marine Estate.

This report is available on The Crown Estate website at: www.thecrownestate.co.uk

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Disclaimer

The opinions expressed in this report are entirely those of the authors and do not necessarily reflect

the view of The Crown Estate, and The Crown Estate is not liable for the accuracy of the information

provided or responsible for any use of the content.

Suggested Citation

Craig J. and Priede, I.G. 2012.

‘Bioluminescence – a source of marine energy?’

The Crown Estate, 27 pages. ISBN: 978-1-906410-32-2.

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Contents

Summary (i)

1. Introduction 1

2. Marine bioluminescence in the British Isles and adjacent seas 2

3. Energy extraction options 4

4. Bioluminescent energy calculations 7

5. Energy extraction proposals 9

5.1. Algae 9

5.2. Jellyfish 12

5.3. Bacteria 15

5.4. Leisure and tourism 16

6. Conclusions 17

References 19

Summary

This study investigates the potential for extraction of energy from bioluminescent marine organisms.

Bioluminescence species occurring around the British Isles and adjacent waters are listed. The report

considers a range of energy extraction options. The bioluminescent characteristics of various marine

animals, algae and bacteria are considered and the potential energy output calculated. Scenarios are

outlined in which energy harnessed from the production of bioluminescent light could be integrated

with existing and emerging commercial activities.

The most promising technology is the culture of planktonic algae (dinoflagellates) using methods

already developed for the production of algal biofuel. However, the energy available from

bioluminescence is less than 1% of that produced by the biofuel. We estimate that a plant extending

over 7 hectares would be necessary to sustain 1 kW of power generation exclusively from

bioluminescence.

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1. Introduction

Bioluminescence refers to light produced by living organisms. It is rare among terrestrial organisms

and virtually absent from freshwater environments. The majority of bioluminescent organisms live in

the marine environment, where they occur from the equator to the polar regions and from the coast

to the deep sea. A wide range of marine organisms are able to bioluminesce, including bacteria,

algae, fish, squid, crustaceans and jellyfish (Herring 1987; Haddock et al. 2010).

Bioluminescent light is produced by a chemical reaction. Although the chemistry is not the same for

all organisms, they do share the same base reaction. In this reaction, a luciferin (protein) reacts with

oxygen in the presence of a catalysing luciferase (enzyme). This produces an oxyluciferin which emits

a photon as it decays from a high energy state to a low energy state. Almost all the energy released

during the reaction is converted into light. In some cases, particularly among jellyfish, the oxygen,

luciferin and luciferase are combined in one molecule, a photoprotein, which emits lights when a

specific ion (such as Ca2+ or Mg2+) is added to the system (Shimomura 2006).

Although bioluminescence ranges in colour from ultra violet (wavelength, λ < 400 nm) to red (λ >

700 nm), most bioluminescence is blue or blue-green (ca. λ 450 – 500 nm), corresponding to the

wavelength of maximum optical transparency of seawater (Figure 1; Widder 2010). The intensity of

bioluminescent flashes largely depends on the emitting organism and ranges over several orders of

magnitude, from 108 to 1013 photons flash-1 (Figure 2; Priede at al. 2008).

In all organisms, except bacteria, light emission is controlled and only emitted in response to a

stimulus. Bioluminescent flashes generally range in duration from 10-1 to 101 seconds. In contrast,

bacteria only produce light when their population reaches a certain density threshold. When this

threshold is reached, bacteria produce a continuous glow. Light is produced by organisms for

defence, to find prey and for communication (Haddock et al. 2010).

Bioluminescence is most frequently used as a defence mechanism. This defence response can be

provoked by mechanical, electrical or chemical stimulation of the organism. The type and strength of

stimulation affects the intensity and duration of the resultant bioluminescent event. This possibly

provides a means of controlling bioluminescent light output and may facilitate its use as a resource.

To date the commercial application of bioluminescence has concentrated on a number of biomedical

and environmental assessment applications. The current major applications of bioluminescence

include using the firefly luminescence system as a method of measuring ATP (adenosine

triphosphate; used as a measure of biological activity); using photoproteins, such as aequorin from a

jellyfish, to detect intracellular Ca2+ (used to regulate various important biological processes), using

Cypridina luciferin within probes for measuring superoxide anions (an indicator of many pathological

conditions) and the use of bioluminescent bacteria to monitor pollution (Shimomura 2006; Girrotti

et al. 2008). There are also reports that during WWII, the Japanese collected large amounts of

bioluminescent ostracods (marine crustaceans), planning to use the material as a source of low

intensity light for use in the jungle, although it appears that none of the material was actually used

(Shimomura 2006).

The aim of this report is to investigate possibilities for harnessing bioluminescence for energy

production, as well as explore alternative avenues of profitable endeavour.

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Figure 1. Mean spectral maximum for bioluminescent animals living in shallow, deep

and benthic (seafloor) marine regions, wavelength in nm (adapted from Widder 2010).

Figure 2. Estimated quantum

emission (photons flash-1 into a

4π steridian sphere) for several

groups of marine bioluminescent

organisms: Copepoda;

Euphasiiacea (Euphaus.);

Amphipoda (Amph.);

Decapoda (De.); Scyphozoa

(Scypp.); Siphonophora (Siph.);

Pyrosoma (Py.); Dinoflagellata

(from Priede et al. 2008; Latz et

al. 2004a).

2. Marine bioluminescence in UK waters

Marine bioluminescence occurs among a wide range of different organisms. A list of the known

bioluminescent species around the British Isles and adjacent seas is shown in Table 1. Coastal species

and species that have been reported within the British coastal region are also indicated.

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Table 1. Bioluminescent marine organisms reported in the British Isles and adjacent seas (inclusion area indicated in light blue on the inserted map). *Coastal species or species that have been reported in coastal areas. (Tett 1971; Herring 1976; Buskey & Swift 1985; Herring 1985; Herring 1987; Widder et al. 1989; Tett 1992; Haddock & Case 1999; Poupin et al. 1999; Haddock et al. 2010; MSBIAS (www.marinespecies.org); MarLIN (www.marlin.ac.uk))

Classification Species Bacteria Vibrio fischeri Vibrio harveyi Vibrio logei Dinoflagellata (Algae) Alexandrium tamarense Ceratium furca Ceratium fusus Ceratium horridum Ceratium tripus Gonyaulax catenata* Gonyaulax polygramma Gonyaulax scrippsae Gonyaulax spinifera Lingulodinium polyedrum Noctiluca scintillans Peridinium divergens* Peridinium ovatum* Peridinium steinii* Polykrikos schwartzii* Prorocentrum micans Protoperidinium bipes Protoperidinium brevipes Protoperidinium cerasus Protoperidinium claudicans Protoperidinium conicoides Protoperidinium conicum Protoperidinium crassipes Protoperidinium curtipes Protoperidinium depressum Protoperidinium divergens Protoperidinium excentricum Protoperidinium leonis Protoperidinium minutum Protoperidinium nudum Protoperidinium oceanicum Protoperidinium ovatum Protoperidinium pallidum Protoperidinium pellucidum Protoperidinium pentagonum Protoperidinium punctulatum Protoperidinium pyriforme Protoperidinium saltans Protoperidinium steinii Protoperidinium subinerme Amphipoda Cyphocaris anonyx Cyphocaris richardi Scina borealis Scina crassicornis

Copepoda Centraugaptilus horridus Euaugaptilus magnus Heterorhabdus norvegicus Heterorhabdus papilliger Heterorhabdus spinifrons Lucicutia flavicornis Lucicutia grandis Metridia longa Metridia lucens* Oncaea conifera Pleuromamma abdominalis Pleuromamma borealis Pleuromamma gracilis Pleuromamma robusta Pleuromamma xiphias Decapoda Acanthephyra pelagica Acanthephyra purpurea Pasiphaea tarda Sergestes arcticus Sergia robustus (robusta) Systellaspis debilis Euphausiidae Meganyctiphanes norvegica* Nyctiphanes couchii Stylocheiron longicorne Stylocheiron maximum Thysanoessa inermis Thysanoessa longicaudata Thysanoessa raschii* Ostracoda Conchoecia spinirostris Ophioroid Amphipholis squamata* Amphiura filiformis* Polychaeteae Tomopteris helgolandica Tomopteris septentrionalis Chaetognath Sagitta elegans Sagitta tasmanica Appendicularian Oikopleura dioica* Thaliacian Cyclosalpa bakeri Pyrosoma atlanticum Hydroida (Medusae) Aequorea forskalea* Aequorea macrodactyla Aequorea vitrina Campanularia sp.* Clytia hemisphaerica* Eutonina indicans Halicreas minimum Halopsis ocellata Lizzia sp.* Obelia sp.* Octophialucium funerarium Tima bairdi* Leptomedusae (Medusae) Cosmetira pilosella* Mitrocomella polydiademata*

Trachylina (Medusae) Aegina citrea Aeginura grimaldii Solmissus incise Coronatae (Medusae) Atolla parva Atolla vanhoeffeni Atolla wyvillei Nausithoe atlantica Nausithoe globifera Paraphyllina ransoni Periphylla periphylla Semaeostomeae (Medusae) Chrysaora hysoscella Pelagia noctiluca* Ctenophora Beroe cucumis* Bolinopsis infundibulum* Siphonophora Abylopsis tetragona Chuniphyes multidentata Diphyes dispar Hippopodius hippopus Nanomia cara Praya dubia Rosacea plicata Vogtia glabra Vogtia serrata Bivalvia Pholas dactylus * Cephalopoda Bathyteuthis abyssicola Brachioteuthis bowmani Brachioteuthis picta Brachioteuthis riisei Galiteuthis armata Gonatus steenstrupi Heteroteuthis dispar * Histioteuthis bonnellii Histioteuthis reversa Loligo vulgaris* Mastigoteuthis schmidti Octopoteuthis sicula Ommastrephes bartramii Ommastrephes pteropus Onychoteuthis banksii Sepiola affinis Sepiola atlantica* Sepiola rondeletii* Spirula spirula * Teuthowenia megalops Elasmobranchii Etmopterus princeps Etmopterus spinax Osteichthyes Argyropelecus hemigymnus Argyropelecus olfersii Epigonus telescopes Himantolocphus groenlandicus Malacocephalus laevis* Maurolicus muelleri Nezumia aequalis Sternoptyx diaphana Xenodermichthys copei

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3. Energy extraction options

There are a number of options available to exploit the energy resources of marine organisms. The

most common and direct use of marine energy reserves is as nutrition for both humans and animals

(Fig 3A). However, in terms of a source of extractable energy, the simplest option is to directly

combust the organisms to produce heat energy (Fig. 3B: Crude fuel). All organisms are primarily

composed of water, carbohydrates, proteins and lipids. The latter three of these substances have

calorific values of 17.2 kJ g-1, 23.7 kJ g-1 and 36.1 kJ g-1, respectively (Lucas 1993). Table 2 shows the

intrinsic calorific content of range of marine organisms. The calorific content of the ash free dry

weight of these organisms ranges over an order of magnitude, from 5.50 x 103 kJ kg-1 for marine

bacteria to 3.05 x 104 kJ kg-1 for the copepod Metridia longa.

Figure 3. Schematic of energy extraction options from the biomass of marine

species, including bioluminescent organims. A: Direct nutrition, B: Crude fuel,

C: Refined fuel, D: Bioluminescent light source, E: Bioluminescent energy source.

The combustion process produces heat energy which can then, if required, be converted into

electrical energy. However, a number of energy costs are involved in this process, in particular, the

evaporation of the water content from the organism (see Box 1). In addition to this, the efficiency of

the heat to electrical energy conversion is limited by the theoretical Carnot Limit1. In reality this

theoretical efficiency value is never reached, although recent studies claim that up to 90% of the

Carnot Limit may be achievable with technological advancements (web.mit.edu; Wu et al. 2009).

Thus the final net energy output will be a fraction of the intrinsic energy content.

1 A theoretical value which sets an absolute limit on the efficiency with which heat energy can be turned into

useful work.

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An alternative method to exploit the energy reserves of marine species is the targeted extraction of

high energy and/or high value products, such as oil (Fig. 3C). The extraction of oil from algae for

biofuel has been in consideration for a number of years and extensive research is being conducted

into its viability as a commercial enterprise. This option is discussed in more detail in section 5.1.

A further option for energy exploitation of marine species is to use the light produced by

bioluminescent species. In this case, the light may either be used directly as a source of illumination

(Fig. 3D), or converted into electricity (Fig. 3E). To assess the potential energy production from

bioluminescent light, the flash characteristics of different species are considered in section 4.

Table 2. Calorific, water and lipid (oil) content of a range of marine groups and species. These values have

been reported in the literature, but it should be noted that the biochemical composition of species varies

with the age and sex of the organisms as well as the environment in which they grow (Raymont et al. 1971;

Finlay & Uhlig 1981; Croxall & Prince 1982; Larson 1986; Ikeda & Skjoldal 1989; Simon & Azam 1989; Davies

1993; Ikeda & Hirakawa 1998; Mansour et al. 1999; Kim et al. 2010; Nurnadia et al. 2011). *Mean value

calculated from a range of reported values.

Group Species

Calorific content (kJ kg

-1 of ash free dry weight)

Water content (% wet weight)

Lipid (oil) content

(% dry weight)

Marine bacteria 5.50x103

46 - 82 (% volume)

Dinoflagellates 2.29x104 12.6 *

Noctiluca spp. 6.60x103

Medusae 1.89x104 95 - 97 3.4 *

Ctenophores 1.68x104 95 - 97

Beroe cucumis 1.92x104 96

Bolinopsis infundibulum 1.47x104 0.5 *

Polychaetes 2.36x104 *

Tomopteris helgolandica 2.89x104 * 22

Copepods 2.31x104 64 - 82 20.1

Metridia longa 3.05x104 * 77

Euphausiids 2.33x104 68 - 82 15.5

Thanoassa spp. 2.27x104 68 - 82 14.7

Meganyctiphanes norvegica 2.78x104 * 75 - 80 17.7 *

Amphipods 1.97x104 * 78 - 83 9.5

Chaetognaths 2.27x104 * 89 6.5

Fish 2.42x104 60 - 82

Squid 2.34x104 74 - 84 19.4

Appendicularia 95 - 97 6.5 *

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Box 1. Some of the issues that surround bio-energy can be understood by considering the case of a draught horse. The standard value for 1 horsepower is 750 W (Watts) based on the original estimate of the continuous working capacity of a brewery horse by James Watt in 1782. The direct energy content of the carcase of a mammal is approximately 7000 kJ kg-1 live weight. Assuming the horse weighs 750 kg this represents a chemical energy store of 5,250,000 kJ which could theoretically be released if the carcase was combusted in a furnace. If we compare this with keeping the horse alive and working it at 750 W, it would take 81 days, working 24h per day to realise the same amount of energy. Working a horse continuously for 24 h a day is of course not feasible and also 750 W is a rather optimistic energy output for a horse. The horse would also have to be fed at the equivalent of over 750 W to maintain it, assuming 30% efficiency ca. 220 MJ day-1. In the case of bioluminescence, having grown some algae (or other bioluminescent organism) we can chose to either extract the stored chemical energy by combustion or keep the algae alive and repeatedly extract energy by stimulating bioluminescence. The algae would need a supply of nutrients to sustain this in the same way as the horse has to be fed. However, data in the literature giving the chemical energy content of various animals, crops and foods (which is useful for dietary consumption estimates) ignores the practical problem of removing the water content before combustion takes place. The latent heat of evaporation of water is: 2260 kJ kg-1. To this must be added the energy required to heat the water to boiling point (100°C) from room temperature (20°C) = 4.18 × 80 = 334 kJ kg-1 So total cost of water removal is 2260 + 334 = 2594 kJ kg-1. For the horse carcase, assuming 70 % water content (525 kg water) this would be an energy loss of 1,362,060 kJ reducing the combustion energy available to 3,887,940 kJ In most marine biomass the water content is somewhat higher e.g. using the same figures but with 85% water content. The energy content of the carcase decreases to: 2,625,000 kJ The cost of water removal increased to: 1,653,930 kJ The net energy available is then 971,070 kJ This indicates that for most wet marine biomass direct combustion is not likely to be profitable owing to the high cost of drying. For traditional biomass, e.g. wood and peat, the water content is much lower and the drying is done for “free” by standing the material for a year or more, essentially dried by solar energy which is not accounted for. Oil can be extracted without drying by disrupting cells, mechanically, chemically or by heat and allowing oil to float to the surface.

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4. Bioluminescent energy calculations

The intensity, duration and spectral characteristics of bioluminescent emissions have been measured

for relatively few species. However, table 3 shows the average bioluminescent light characteristics of

several species, covering a range of different animal groups and dinoflagellates (algae).

Bioluminescent bacteria are not listed in table 3 as they generate a continuous glow, not discrete

flashes of light (see Section 5.3 for more details on bacterial bioluminescence).

The energy of a single flash, from the organisms listed, varies from 10-12 to 10-9 kJ flash-1. Based on

these values, a calculation has also been made to show the number of individual flashes that would

be required to create 1 kW of light, which ranges in magnitude from 108 to 1011.

Bioluminescent flashes may be stimulated multiple times in a single organism. Stimulation may be

provided by mechanical, electrical or chemical means. It is important to note, however, that after an

initial bioluminescent flash, subsequent flashes can be of lower intensity, as the bioluminescent

potential of the organism becomes exhausted. The full bioluminescent potential is restored given a

sufficient period of time. This recovery time will vary between species and on the initial state of

exhaustion. In the case of dinoflagellates, for example, studies have shown that recovery can occur

in 0.5 – 6 hours (Widder & Case 1981).

To convert bioluminescent light into electrical energy, photovoltaic cells would be required. Thus the

final energy output from the conversion of light to electricity would be dependent on the efficiency

of the photovoltaic cell. The efficiency of photovoltaic cells used for the conversion of solar light to

electrical energy tend to be in the region of 10 – 15 % at a light intensity of 1 sun, although recent

advances in the use of different materials report efficiencies up to 42 % (Green et al. 2011). Such

high efficiencies are reported for high light intensities, > 1 sun. The average sunlight intensity in the

U.K. is about 100 W m-2 (Mackay 2008); several orders of magnitude brighter than bioluminescent

light. At lower light intensities the power efficiency of silicon and organic photovoltaic cells are less

efficient (Reich et al. 2009, Steim et al. 2011). However, demand for photovoltaic cells for indoor use

has lead to the development of techniques to improve their efficiency at lower light levels (Steim et

al. 2011).

Photovoltaic cells designed to convert sunlight into electrical energy are designed to be able to

absorb light from a wide spectral window to respond to the broadest possible span of the solar

spectrum. This is achieved using a stack of photovoltaic cells each with peak sensitivity in different

regions of the solar spectrum (Forrest 2005). Most bioluminescent light emission is blue or blue

green (ca. λ 450 - 500 nm) and of narrow bandwidth (λ 26 – 100 nm) (Widder et al. 1983), so

photovoltaic cells sensitive to the blue bandwidth (e.g. Zhang et al. 2010) would be most suitable for

the conversion of bioluminescent light into electrical energy.

In the future, there may be possibilities of producing a higher bioluminescent light yield through

genetic modification of organisms. Non-bioluminescent bacteria can be genetically engineered to

bioluminesce (Sagi et al. 2003; Girotti et al. 2008) and a successful attempt was made to increase the

light output from an extracted luciferase (Fujii et al. 2007). However, no studies to date can be found

demonstrating an enhanced light yield from a live multicellular animal. Thus this approach would

require extensive research and development.

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Table 3. Light and energy output estimates for bioluminescent emissions from a range of marine species.

(Nicol 1958; Clark et al. 1962; Swift et al. 1977; Lapota & Losee 1984; Latz et al. 1990; Bowlby et al. 1990;

Bowlby & Case 1991; Latz et al. 2004a; Priede et al. 2008)

Classification Species Quantum emission

(photons flash−1

)

Peak wavelength

(nm) (*assumed)

Average duration

(s)

Energy of single flash (J flash

−1)

Flashes required for

1 kW

Copepoda

Gaussia princeps 1.80x1011

480* 8.7 7.45x10-8

1.17x1011

Pleuromamma xiphias

1.80x1010

480* 0.4 7.45x10-9

5.07x1010

Metridia lucens 3.44x1012

480 6.7 1.42x10-6

4.70x109

Euphausiiacea

Meganictyphanes norvegica

7.55x1010

480 13 3.12x10-8

4.16x1011

Euphausia eximia 1.00x1010

480*

4.14x10-9

Nyctiphanes simplex

1.00x1011

480*

4.14x10-8

Amphipoda

Cyphocaris faurei 5.70x1010

480*

2.36x10-8

Scina crassicornis 3.10x1010

480*

1.28x10-8

Decapoda Acanthephyra pelagica

2.73x1012

490 3.5 1.11x10-6

3.16x109

Scyphozoa

Periphylla periphylla

3.50x1010

465 2.4 1.50x10-8

1.61x1011

Atolla wyvillei 1.20x1011

470 2 5.07x10-8

3.94x1010

Siphonophora

Vogtia glabra 1.40x1011

470 4 5.92x10-8

6.76x1010

Vogtia spinosa 3.20x1011

470 6.5 1.35x10-7

4.81x1010

Pyrosoma Pyrosoma atlanticum

2.30x1013

493 59 9.27x10-6

6.37x109

Dinoflagellata

Ceratium fusus 1.10x109 474 0.2 4.61x10

-10 5.18x10

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Ceratocorys horrida

9.20x109 474 0.2 3.86x10

-9 4.77x10

10

Lingulodinium polyedrum

1.90x108 474 0.1 7.96x10

-11 1.57x10

12

Pyrocystis fusiformis

6.90x1011

472 0.2 2.90x10-7

7.23x108

In terms of the conversion of bioluminescent light into electricity, the current limited efficiency of

photovoltaic cells, combined with the low light intensity of the bioluminescent emissions (relative to

solar light), may render this option uneconomic as a stand-alone technology. However, it may be

feasible to integrate the harvesting of light with another energy extraction process. In this report we

consider the integration of bioluminescent energy extraction with various existing and emerging

commercial activities.

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5. Energy extraction proposals

5.1 Algae

In recent years there has been a lot of interest in the use of algae to produce biofuel. There are

several benefits that are offered by this biofuel production option compared to other methods

which extract oil from land crops. One of the principal advantages is the short growth cycle of algae

which increases the potential productivity by several orders of magnitude per unit area compared to

terrestrial plant options. In addition to this, algae can be produced on non-arable land so does not

compete with food production (Turner et al. 2011). However, challenges do remain before algal

biodiesel can become a viable option. One of the main challenges to making this process viable is the

high energy input required to dry the algae to extract the oil. The drying process requires large

amounts of fossil fuel derived energy, currently rendering algal biofuel uneconomic and

unsustainable (Sander & Murphy 2010). However, further research and development is currently

underway. In the U.K., the Carbon Trust Challenge has invested in the development of a sustainable,

cost-effective biofuel from algae (www.carbontrust.co.uk). Within the E.U., the European Industrial

Bioenergy Initiative (EIBI) supports demonstration and flagship projects for the production of

bioenergy that is not yet commercially available (www.biofuelstp.eu).

There are two ways that the economic viability of algal biofuel production can be augmented: either

by extracting additional high value co-products from the cultured algae; or by increasing the energy

output of the process. Various high value co-products are currently being considered, including

animal and fish feed, food and food additives, chemical feedstocks, health and beauty products, as

well as pharmaceuticals (Dyer-Smith & Allen 2011). In addition to these revenue generating

products, processes that increase the energetic yield during the production of algae will also

improve the net energy balance of algal biofuel production. Harvesting bioluminescent light from

algae during the incubation period has the potential to serve as an additional source of energy

output. The combination of these energy extraction options is illustrated by figure 4, in which

bioluminescent light is stimulated within the biomass during incubation, prior to the extraction of

oil.

Figure 4. Schematic of option for extraction of bioluminescent light and refined

fuel from the biomass of marine algae.

The only class of algae that includes bioluminescent species is Dinophyceae; the dinoflagellates.

Although there are over 3000 strains of algae that are being considered for their suitability for

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biofuel production, the most appropriates classes are the green algae and the diatoms (Sheehan et

al. 1998). These algal groups have been considered most appropriate for the quantity and type of

lipids they produce. However, the quality and quantity of lipid content can be enhanced through

genetic improvement and modification (Lundquist et al. 2010; Radakovits et al. 2010; Singh et al.

2011). Although the authors of the present report find no evidence for current investigations into

the production of a suitable bioluminescent dinoflagellate species or a strain genetically engineered

to bioluminesce, this remains a possible avenue for future research.

Dinoflagellates require mechanical stimulation above a certain threshold to produce

bioluminescence (Latz et al. 1994; Latz et al. 2004a,b; Cussatlegras & Le Gal 2005). The threshold

varies between species, but in general, an accelerating flow with shear force (≥ 0.6 N m-2) is required

to maximise the bioluminescent output (Blaser et al. 2002; Cussatlegras & Le Gal 2005). Within an

algal production plant the mechanical stimulation could be generated by a purpose built system.

However, mechanical stirring is a necessary component of the algal cultivation process and

potentially it would be possible to integrate the process of stirring and stimulation into one unit.

Two of the systems in consideration for the cultivation of photosynthetic algae are raceway ponds

(open systems) and photobioreactors (PBRs; closed systems) (Chisti 2007). In PBRs the algae is

contained within transparent structures enabling greater control over the environmental conditions

and reducing the risk of contamination. During the algal biofuel production process turbulence is

created in both the open and closed systems which could be used as a site of bioluminescence

stimulation. In the open system, a unit is required to mix and circulate the algal broth. In the closed

system, turbulence is required at two points in the process; within the growth chambers to prevent

sedimentation and also at the degassing zone to extract the build up of oxygen within the algal

broth. Where turbulence is created, a bioluminescent species would be stimulated to produce

bioluminescence.

Photosynthetic algae2 require a day and night light regime for optimum growth. Daylight is known to

inhibit the production of bioluminescent emissions in photosynthetic dinoflagellates (Hamman et al.

1981; Buskey et al. 1992). As a result of this photoinhibition of bioluminescent emissions during the

light cycle, bioluminescent light could only be harvested during the dark cycle. Using production

values that have been previously estimated for the algal biofuel process, it is possible to estimate

the potential bioluminescent light output from a hypothetical production plant. Table 4 shows the

estimated energetic output from a range of dinoflagellate species. The energetic output per unit

weight is species dependent, ranging from 6.37 x 10-4 to 2.32 kJ kg-1. Assuming an algal production

plant can produce an algal biomass of 22 g m-2 day-1 (following the estimate of Lundquist et al.

(2010)) the potential energetic output from the stimulation of bioluminescent flashes depends on

the number of times a day each cell is stimulated. Although bioluminescence may be stimulated

repeatedly in a single organism, the bioluminescent potential of the organism becomes exhausted

often resulting in subsequent flashes of lower intensity. The full recovery time varies between

species, but can be assumed to be 0.5 – 6 hours in dinoflagellates (Widder & Case 1981). Thus, if

each cell were stimulated twice a day the energetic output would range from 2.80 x 10-5 kJ m-2 day-1

for the species Lingulodinium polyedrum to 1.02 x 10-1 kJ m-2 day-1 for the species Pyrocystis

fusiformis. Increasing the rate of stimulation of each cell to from 2 to 24 times a day would increase

2 Photosynthetic algae convert sunlight into chemical energy, including oil.

11

the potential energetic output of each species by just over an order of magnitude (Table 4),

assuming full recovery of bioluminescent potential between flashes.

Table 4. Estimated energetic output per unit weight (kJ kg-1

) and the daily energetic output per unit area assuming a theoretical algal production scenario (kJ m

-2 day

-1) for a range of dinoflagellate species.

Dinoflagellate species

Energy output

a

(kJ kg-1

)

Energy output a,b,c

(kJ m-2

day-1

)

assuming stimulation of each cell twice per day

Energy output a,b,d

(kJ m-2

day-1

)

assuming stimulation of each cell 24 times per day

Ceratium fusus 3.69x10-3

1.62x10-4

1.95x10-3

Ceratocorys horrida 3.08x10-2

1.36x10-3

1.63x10-2

Lingulodinium polyedrum

6.37x10-4

2.80x10-5

3.36x10-4

Pyrocystis fusiformis 2.32 1.02x10-1

1.23

a

Assuming a wet weight of 1.6x10-7

g per dinoflagellate cell b Assuming a total biomass productivity of 22g m

-2 day

-1 (Lundquist et al. 2010).

c Assuming a 6 h recovery period of bioluminescent potential for each dinoflagellate cell, with light harvested

during a 12 h dark period of the growth cycle. d Assuming a 0.5 h recovery period of bioluminescent potential for each dinoflagellate cell, with light harvested

during a 12 h dark period of the growth cycle.

Converting the bioluminescent light produced during the algal production process into electrical

energy would increase the net energy balance of the algal biofuel production process. As an

indication of the significance of the bioluminescent energy contribution, the energetic output from

the dinoflagellate bioluminescence can be compared to the energetic output from the oil content of

the algae. Lundquist et al. (2010) estimate that the daily oil production from this algal production

plant would be 5.5 x 10-3 l m-2 day-1 (equivalent to 20000 l hectare-1 year-1). The energy content of oil

is 33 MJ l-1, giving a daily energy output of 180 kJ m-2 day-1. This calculation does not take into

account any of the energetic costs of processing or drying the algae it simply provides the potential

energy output of the oil content. Therefore the potential energy output from the bioluminescence of

the dinoflagellate Pyrocystis fusiformis (1.23 kJ m-2 day-1) represents about 1% of the energy

potential of its oil content. If this output from bioluminescence can be achieved, it would require an

area of 70244 m2 or over 7 hectares to generate 1kW of power.

The light and temperature regimes in the U.K. are not considered optimum for the production of

photosynthetically derived algal biofuel. The Carbon Trust Challenge aimed to undertake the

production of algal biofuel in equatorial regions where temperatures are higher and light regimes

are stable throughout the year. Thus, the production of algae in pond raceways and PBRs may not be

suitable to the U.K. climate.

However, alternative systems have also been researched and developed which use different

production techniques or create alternative fuels to oil (Mascarelli 2009; Savage 2011). One

technique which has been developed in the U.S. by the company Solazyme uses heterotrophic3 algae

which grow in the dark by consuming sugars (Mascarelli 2009; solazyme.com). In 2010, Solazyme

3 An organism that cannot synthesize its own food and is dependent on complex organic substances for

nutrition.

12

supplied the U.S. Navy with 80,000 l of algal derived diesel and jetfuel using this technique. Other

alternatives include the harvesting of ethanol, which is excreted by some types of algae, as

developed by the U.S. company Algenol Biofuels (www.algenol.com). Other companies that are

involved in the production of algal biofuel include Petrosun, Green Star, SGI, Solix Biofuels, Sapphire

Energy, BioFuel Systems SL, Petro Algae, and HR Biopetroleum. The possibility exists that

bioluminescent light harvesting could be applied to any of these techniques.

5.2 Jellyfish

Bioluminescence in common among jellyfish4 and around the British Isles and adjacent waters there

are at least 37 known bioluminescent species of medusae, nine bioluminescent species of

siphonophore and two bioluminescent species of ctenophore (Table 2). In this section, the possibility

of combining the extraction of bioluminescent energy from jellyfish with other commercial

applications is explored.

Jellyfish play a natural and important part of marine ecosystems. However, there is concern that

jellyfish numbers may be increasing worldwide and that jellyfish blooms are becoming more

frequent, although confirmation of such trends is hindered by the absence of baseline data (Mills

2001; Pauly 2009; Richardson et al. 2009; Purcell 2012; www.bbc.co.uk/news). A number of

anthropogenic factors are believed to contribute to increased jellyfish numbers. Overfishing reduces

the numbers of competitors and predators of jellyfish, allowing their population to increase (Parsons

& Lalli 2002; Richardson et al. 2009). Eutrophication (increased concentrations of nutrients from

fertiliser runoff and sewage) encourages the development of algal blooms which can create

favourable conditions for the proliferation of jellyfish (Purcell 2001). Additionally, warming of the sea

surface as a result of climate change may also increase jellyfish numbers (Parsons & Lalli 2002;

Purcell et al. 2007; Gibbons & Richardson 2008; Lynam et al. 2010). An EU funded project called The

EcoJel Project (www.jellyfish.ie) was established in 2007 to assess the opportunities and detrimental

effects of jellyfish in the Irish Sea.

One of the major issues of increasing numbers of jellyfish is their predation of vulnerable life stages

of fish (fish eggs and larvae) (Hay 1990; Paurcell & Arai 2001; Lynam et al. 2005). It is hypothesized

that increasing jellyfish populations have the potential to displace fish communities (Purcell 2007;

Richardson et al. 2009). This would shift the ecosystem to a more gelatinous one, having a

detrimental effect on fish stocks. There is also evidence that jellyfish blooms cause high mortality of

farmed fish (Bämsted et al. 1998; Doyle et al. 2008; Ferguson et al. 2010; Baxter et al. 2011) and

have been responsible for blocking the cooling systems of nuclear reactors (www.bbc.co.uk/news).

Within UK waters there is some evidence to suggest that jellyfish numbers are increasing in the

North Sea (Attrill et al. 2007) and the Irish Sea (Lynam et al. 2011). If jellyfish numbers and blooms

are increasing in UK waters it would be pertinent to take advantage of this resource as part of any

management approach.

4 Free swimming gelatinous animals that include medusae, siphonophores and ctenophores.

13

Jellyfish could be fished or cultured to exploit their bioluminescent properties. The harvest of

jellyfish could also be combined with other commercial uses of jellyfish (Fig. 5). Table 4 provides an

estimate of the energy and power output of a range of jellyfish species per unit weight. Although the

energy produced per flash is of the same order of magnitude (10-8 J; Table 3) for each of the species

listed in Table 5, the energetic output per unit weight varies over two orders of magnitude, from

10-10 to 10-8 kJ kg-1, as a result of differences in the average weight of individual jellyfish.

Figure 5. Schematic of option for extraction of bioluminescent light and

various valuable products from jellyfish.

Table 5. Energy output per unit weight for a range of jellyfish (Lancraft et al. 1991; Lucas 2009; Lebrato & Jones 2009)

Jellyfish species Average wet weight (g) Energy output a

(kJ kg-1

) Power output a (kW kg

-1)

Periphylla periphylla 6 2.72x10-9

1.13x10-9

Atolla wyvillei 64 7.98x10-10

3.99x10-10

Vogtia glabra 1 5.61x10-8

1.40x10-8

a Assuming stimulation of each individual once

Jellyfish are currently used for food, clinical and industrial applications. In Asia, semi-dried jellyfish

represents a multi-million dollar seafood business (Hsieh et al. 2001; Omori & Nakano 2001; You et

al. 2007). Edible jellyfish are supplied by both aquaculture techniques (You et al. 2007) and jellyfish

fisheries (Omori & Nakano 2001; Nishikawa et al 2008). In some areas of China, jellyfish (Rhopilema

esculentum) aquaculture has now replaced traditional fish and shrimp aquaculture (You et al. 2007).

Most of the world’s jellyfish fisheries are based in Asia, where demand has been growing since the

1970s (Kingsford et al. 2000; Omori & Nakano 2001). In response to this demand, jellyfish fisheries

are expanding worldwide. The US has started harvesting the Cannonball jellyfish (Stomolophus

meleagris) for export to the food markets of Asia where there is high demand. Cannonball jellyfish

are very abundant in coastal US waters and are often problematic for net based fisheries. The

jellyfish can crush the target species or even damage nets as a result of their weight and bulk.

Harvesting these jellyfish provides economic and ecological benefits (Hsieh et al. 2001). In addition

14

to those of the U.S., jellyfish exports are reported from Australia, India (Hsieh et al. 2001) Namibia

and the U.K.5 (You et al. 2007).

Omori & Nakano (2001) list the most important edible jellyfish species (Table 6). None of the listed

species are bioluminescent. However, one of them, the Dustbin-lid jellyfish (also known as the Barrel

jellyfish; Rhizostoma pulmo; synonym R. octopus), is present on the western and southern coasts of

Britain (Sabatini 2004) and is considered a bloom forming species (Lilley et al. 2009). Although this

particular species is not of interest from the perspective of bioluminescence applications, it may

have potential for future commercial exploitation in British waters as an export to the Asian food

market.

Table 6. Identified species of edible jellyfish (Omori & Nakano 2001)

Cephea cephea (Forskål, 1775) Catostylus mosaicus (Quoy & Gaimard, 1824) Crambione mastigophora Maas 1903 Crambionella orsisi (Vanhöffen, 1888) Lobonema smithii Mayer, 1910 Lobonemoides gracilis Light, 1914 Rhizostoma pulmo (Macri, 1778) Rhopilema esculentum Kishinouye, 1891 Rhopilema hispidum (Vanhöffen, 1888) Neopilema nomurai Kishinouye, 1922 Stomolophus meleagris L. Agassiz, 1862

Jellyfish are also exploited for their medicinal values. Although there is little scientific evidence for

the purported effectiveness of jellyfish in weight loss, softening skin, treating burns, arthritis,

fatigue, hypertension, back pain and ulcers; it remains popular in the Asian medicinal market.

However, there have been some recent reports in the scientific literature showing preliminary

positive effects of jellyfish collagen extracts on arthritis (Hsieh et al. 2001; Ohta et al. 2009), fatigue

(Ding et al. 2011) and hypertension (Morinaga et al. 2010). In the West, collagen extracts for

pharmaceutical and nutritional applications are generally obtained from pig, bovine and poultry

sources, although consideration has been given to the benefits of collagen extraction from jellyfish

(Nagai et al. 2000; Addad et al. 2011; Gomez-Guillen 2011).

Pelagia noctiluca is a bioluminescent jellyfish species that has been found to be widespread in Irish

and UK waters (Russell 1970; Doyle et al. 2008; Bastian et al. 2011). It is not an edible species but

there has been some investigation into the extraction of its collagen. However, the concentration of

collagen in the body tissues of P. noctiluca is low compared to the Dustbin-lid jellyfish (R. Pulmo)

(Addad et al. 2011) and it may be unlikely to provide an efficient collagen source.

The photoproteins of several jellyfish species have been used for pharmaceutical applications. These

photoproteins are triggered to luminesce by the addition of the calcium (Ca2+). This property has

5 N.B. The authors of this present report find no further evidence of U.K. exports of jellyfish to the Asian food

market other than the reference in You et al. (2007).

15

been harnessed as a technique to measure intracellular calcium for a wide range of medical research

applications. More than 25 bioluminescent species are known to use Ca2+ regulated photoproteins,

although to date only seven photoproteins have been isolated and characterised (Vysotski et al.

2006). Thus there is scope for the discovery of photoproteins that perform better than those

currently employed. However, once selected for pharmaceutical use, the photoproteins are cloned

and synthetically produced, rather than extracted from the original species (Markova et al. 2002;

Vysotski et al. 2006; Inouye 2008). Consequently, there is likely to be little scope for commercial

exploitation of jellyfish for extraction of their photoproteins.

5.3 Bacteria

Bioluminescent bacteria are currently used for monitoring environmental pollutants (reviewed in

Girotti et al. 2008). However, there are investigations into potential future uses of bioluminescent

bacteria.

Bioluminescent bacteria produce a continuous glow once a certain cell density is reached. As no

other stimulus is required to produce the light, one possible application of bacterial bioluminescence

is as a source of illumination (Fig. 6). The electronics company Philips is currently exploring the

possibility for bioluminescent home lighting in an experimental project called the Microbial Home

System. Part of this project includes a ‘Bio-light’ design (www.design.philips.com), consisting of glass

cells containing live bacterial cultures that emit soft green bioluminescent light in the home (Figure

7). Bioluminescent bacteria glow continuously without the need for any kind of external stimulation

(electrical, mechanical, chemical or photic) making it suitable to such an application. The intensity of

light production is dependent on the species as well as the bacterial cell density. In optimum

conditions the light output of several bioluminescent bacteria species are indicated in Table 7. At a

cell density of 250 x 106 cells ml-1, the brightest bacterial strain of those listed in Table 6, Vibrio

harveyi, would produce 2.03 x 10-7 Watts l-1. This is equivalent to a daily energy output of

1.75 x 10-5 kJ l-1 day-1.

Figure 6. Schematic of option for use of bioluminescent bacteria as a light source.

Although the intensity of light emitted by the Philips prototype is currently deemed insufficient to

act as a primary source of illumination in the home, other applications for bioluminescent bacterial

light have been suggested. These options include night-time road markings (e.g. bioluminescent

plants that indicate where the edge of the road is), warning strips on flights of stairs, kerbsides etc,

informational markings in low-light settings (eg. theatres, cinemas, nightclubs), and new genres of

atmospheric interior lighting with, for example, possible therapeutic and mood-enhancing effects.

16

Figure 7. Prototype of the bioluminescent bacterial lighting design from Philip’s Bio-light

project.

Table 7. Estimated power output from several species of bioluminescent marine bacteria (Bourgois et al. 2001).

Bacteria species

Light output

a

(Photons s-1

cell

-1)

Peak wavelength

(nm)

Power per cell

(W cell-1

)

Cells required for

1 kW

Power per litre

b

(W l-1

)

Daily energetic

output (kJ l

-1 day

-1)

Vibrio harveyi 2 490 8.11x10-19

1.23x1021

2.03x10-7

1.75x10-5

Vibrio fischeri 0.6 490 2.43x10-19

4.11x1021

6.08x10-8

5.25x10-6

Photobacterium phosphoreum

0.5 490 4.93x10-19

4.93x1021

5.07x10-8

4.38x10-6

a Assuming steady state conditions with adequate oxygen supply

b Assuming a cell density of 250 x 10

6 cells ml

-1, indicated as an optimum density for bioluminescent output

(Bourgois et al. 2001).

5.4 Leisure and tourism

As a final note, it is worth mentioning the high degree of public interest in bioluminescent creatures.

Various bioluminescent fantasy forms have featured in the blockbuster films ‘Avatar’ and ‘Monsters’

as well as various video games. In the U.S., the American Museum of Natural History in New York has

announced a new exhibition to run throughout 2012 entitled ‘Creatures of light: Nature’s

bioluminescence’ (www.amnh.org). In addition to this, areas such as Bioluminescent Bay in Puerto

Rico attract huge numbers of tourists to kayak through the glowing waters, rich in bioluminescent

dinoflagellates (www.tripadvisor.co.uk).

In the U.K. there are reports of naturally occurring dinoflagellate bioluminescence in coastal waters

in the Moray Firth (personal obs., B. Ruck, Director of Moray First Marine Ltd) and occasionally in

various locations on the west coast of Britain (Staples 1966; Tett 1992). Although the occurrence of

natural bioluminescence in British waters may not be frequent enough to drive a tourist industry,

there may be scope for development of artificial installations. At present there are two U.K. based

aquaria where bioluminescent flashlight fish are displayed (The Deep in Hull and Bristol Blue Reef).

However, there may be potential for expanding the role of bioluminescence within the British leisure

and tourism industry. Future projects could include educational facilities such as exhibitions, or

possibly more entertainment focussed enterprises such as bioluminescent pools.

17

6 Conclusions

This report explores possibilities for the use of bioluminescence as an energy resource. The energy

released per single bioluminescent flash is species dependent and varies over several orders of

magnitude from 10-11 to 10-6 J flash-1 for marine animals and algae (Table 2). Bacteria emit a

continuous glow when the population density exceeds a certain threshold, with power output in the

region of 10-19 W cell-1 (Table 7).

The bioluminescent light generated by marine organisms could be converted into electrical energy.

Photovoltaic cells are used to convert solar light into electrical energy with efficiencies generally in

the region of 10 - 15%. Photovoltaic cells are less efficient at lower light intensities, so the

conversion of bioluminescent light into electrical energy would likely fall below 10 %.

The low light output of bioluminescent species relative to solar light may render the conversion of

bioluminescent light into electrical energy uneconomic as a stand-alone option. However,

alternative scenarios have been considered in this report including the integration of photic energy

extraction with other commercial activities. The potential energy output has been considered for the

bioluminescent emissions of algae (dinoflagellates), jellyfish and bacteria.

Of the dinoflagellate species investigated, Pyrocystis fusiformis generates bioluminescence of

greatest light intensity, producing 2.32 kJ kg-1 if each cell is stimulated to flash once (Table 4).

Bioluminescent flashes may be stimulated multiple times in a single cell, although time is required

for a cell to recover its full bioluminescent potential. The bioluminescent flashes of dinoflagellates

are short in duration, typically 100 – 200 ms (Table 3).

The energetic output per unit weight of the jellyfish species investigated ranged from 7.98 x 10-10 to

5.61 x 10-8 kJ kg-1 (Table 5). The bioluminescent flashes of jellyfish are usually several seconds in

duration (Table 3).

The energetic output per unit weight for dinoflagellates is 4 to 10 orders of magnitude greater than

for jellyfish species (Table 4 & 5). Although the energy per individual flash is generally higher for

jellyfish species than for dinoflagellate species (Fig. 2; Table 2), dinoflagellates are substantially

smaller (ca. 5 x 10-5 m diameter) than jellyfish species (> 10-2 m diameter). This results in a greater

energetic output per unit weight for dinoflagellates than for larger organisms (jellyfish, fish,

crustaceans etc.). The size of the organism is important to consider in terms of the extraction of

bioluminescent energy, as stimulation (mechanical, electrical or chemical) of larger organisms will

require greater use of energy and resources than the stimulation of smaller organisms.

Integrating the harvest of bioluminescent light with algal biofuel production may be possible.

However, at present the algal strains under consideration for use in this process do not include

bioluminescent dinoflagellates. If a bioluminescent strain were to be selected, the potential annual

energy output would largely depend on the quantum light emission of the species, the minimum

recovery time required between consecutive flashes and the total dark period during which flashes

could be elicited. Using values from the dinoflagellate Pyrocystis fusiformis, the daily potential

bioluminescent energy output from a hypothetical algal biofuel production plant would be

1.23 kJ m-2 day-1 (assuming an algal production of 22g m-2 day-1 ; Lundquist et al. 2010). This

energetic output represents about 1% of the energetic content of the potential oil harvest derived

18

from the same quantity of algae. Using optimistic assumptions, a plant extending over more than 7

hectares would be necessary to sustain an output of 1kW from algal generated bioluminescence.

The possibility of integrating the harvest of bioluminescent light from jellyfish with other commercial

activities was also explored. The commercial activities considered were the harvesting of jellyfish for

export to the Asian food market, extracting jellyfish collagen and the extraction of photoproteins.

However, at present no known species of bioluminescent jellyfish that occurs in British waters is

considered edible or suitable for the extraction of collagen, and photoproteins are manufactured

synthetically. In addition to this, the energetic output per unit weight of jellyfish is several orders of

magnitude lower than that of smaller species, such as dinoflagellates.

There are currently investigations into the possibility of utilising the light generated by

bioluminescent bacteria as a direct source of illumination. At present the light intensity of these

systems are low, but bacterial bioluminescence may have potential to serve as a form of low

intensity warning or information lighting.

In addition to options to extract the energy from bioluminescent emissions, these may also be scope

to extend the use of bioluminescence in the British tourism and leisure industry.

19

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