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Patrick Michael Holligan: a short

biography
CAROL ROBINSON* AND NICHOLAS J. P. OWENS
PLYMOUTH MARINE LABORATORY, PROSPECT PLACE, WEST HOE, PLYMOUTH PL1

3DH,

UK

*CORRESPONDING AUTHOR: carol.robinson@uea.ac.uk

PRESENT ADDRESS: SCHOOL OF ENVIRONMENTAL SCIENCES, UNIVERSITY OF EAST ANGLIA, NORWICH NR4

7TJ, UK

PRESENT ADDRESS: BRITISH ANTARCTIC SURVEY, NATURAL ENVIRONMENT RESEARCH COUNCIL, HIGH CROSS, MADINGLEY ROAD, CAMBRIDGE CB3

0ET,

UK

Received November 30, 2007; accepted in principle November 30, 2007; accepted for publication December 4, 2007; published online
December 11, 2007
Communicating editor: K.J. Flynn

Our understanding of the role of marine phytoplankton

in the global biogeochemical cycles of carbon and
sulphur, particularly the rates and variability of primary
production and calcification, phytoplankton activity and
ecology in the subsurface chlorophyll maximum, and
the use of satellite remote sensing to detect coccolithophore blooms, has progressed substantially through
research initiated, led and undertaken by Patrick
M. Holligan. The enthusiasm and commitment of
Patricks colleagues for both this special issue and the
symposium on Marine Plankton: From Cells to Ecosystems
held in his honour in April 2008 at Plymouth, UK are
testament to the high esteem in which he is held within
the international scientific community. His expertise as
a scientific leader, teacher and writer has inspired students and collaborators over more than three decades.
In particular, his chairmanship during the early 1990s
of the Planning Committee for the Land-Ocean
Interactions in the Coastal Zone (LOICZ) project led
to the establishment of a new core project within the
International Geosphere-Biosphere Programme (IGBP).
As the first Head of the School of Ocean and Earth
Science (SOES), University of Southampton (1998
2002), Patrick had responsibility for substantial expansion of the School, including 13 new academic appointments. As one of the first UK Principal Investigators to
propose an open-ocean latitudinal time-series study, he
helped initiate the development of the Atlantic
Meridional Transect (AMT) programme (www.amt-uk.
org), and as the President of the Council of the Sir
Alister Hardy Foundation for Ocean Science
(SAHFOS), he guided the transition to a new

Directorship of the Continuous Plankton Recorder

programme.
Patrick gained an MA in Botany from Christs
College, Cambridge in 1966. He then moved to the
Department of Botany at the University of Leeds where
he completed his PhD in 1970 on Carbohydrate
metabolism in the fungus Dendryphiella salina under the
supervision of David Jennings. During the first year of
his PhD, Patrick assisted Ed Drew in some research on
acclimations by seaweeds to changes in light intensity
and quality that required summer fieldwork in Malta. It
was here that Patrick practised his self-taught skills in
scuba diving, met John Woods for the first time, acted
as a guinea pig for behavioural research into the effects
of nitrogen narcosis, and developed a serious interest in
the marine environment. While doing his PhD, Patrick
also met and married his wife, Sue. Patrick then undertook a postdoctoral appointment (1969 1973) with
David Lewis in the Department of Botany at the
University of Sheffield to work on the physiology of
higher plants infected by rust fungi.
In 1973, Patrick joined the staff at the Marine
Biological Association (MBA) in Plymouth UK as a
Research Scientist, despite his only marine credentials
being research on a marine fungus and his hobby of
scuba diving. He sought external advice from David
Cushing and Tim Wyatt at Lowestoft and from Jack
Talling at Windermere about developing a new research
programme on marine phytoplankton and then, over
morning coffee, met Robin Pingree, a physicist with an
interest in biology who had come from the Institute of
Oceanographic Sciences (IOS), Wormley, UK to work

doi:10.1093/plankt/fbm106, available online at www.plankt.oxfordjournals.org

# The Author 2007. Published by Oxford University Press. All rights reserved. For permissions, please email: journals.permissions@oxfordjournals.org

JOURNAL OF PLANKTON RESEARCH

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at the MBA. Robin and Patrick formed a working partnership that was to last 10 years and to include about
250 days at sea together (Table I). Together they pioneered the application of continuous measuring techniques and remote sensing to interdisiplinary studies of
shelf seas around the UK, receiving strong support
from colleagues including Gary Mardell, Bob Head,

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Derek Harbour, Dave Griffiths and Phil Pugh (from

IOS) as well as from the Director of the MBA, Eric
Denton.
In 1975, the spring and summer cruises led to the
first detailed descriptions of the development of the
spring phytoplankton bloom and of the biological properties of fronts between well-stratified shelf water and

Table I: Research cruises undertaken by Patrick

1974
1975
1975
1975
1975
1975
1975
1975
1975
1976
1976
1976
1976
1976
1977
1977
1977
1977
1977
1977
1978
1978
1978
1978
1979
1979
1980
1980
1981
1981
1982
1982
1982
1982
1983
1983
1984
1984
1984
1985
1986
1986
1987
1988
1989
1990
1991
1996
1998
1999
2003
2004
2005
2005
2006

3 5 July
9 23 April
9 13 June
17 31 July
19 21 August
7 10 October
13 17 October
3 7 November
8 12 December
13 14 January
9 23 March
20 23 April
20 July 4 August
9 12 August
21 March11 April
30 April 6 May
28 May1 June
11 28 July
16 26 August
30 August 3 September
21 28 April
15 23 June
11 21 July
21 22, 25 27 September
10 23 April
17 30 August
22 April 1 May
29 July 14 August
22 July 2 August
24 August 4 Sept
24 May3 June
18 27 June
20 30 July
2 13 August
16 27 May
July
10 19 June
9 19 July
23 August 2 Sept
10 July 3 August
30 April 15 May
5 22 July
8 June 3 July
9 12 June
19 26 June
27 June 19 July
12 June 3 July
18 April 23 May
9 May16 June
30 July 14 August
25 July 14 August
26 April 2 June
15 July 7 August
12 October 28 Nov
2 14 December

RV Sarsia (Channel grid)

RV Sarsia (W Channel and Celtic Sea)
RV Sarsia (W Channel)
RV Sarsia (W Channel fronts Lundy)
RV Sarsia (E1)
RV Sarsia (Celtic Sea)
RV Sarsia (Celtic Sea)
RV Sarsia (Celtic Sea)
RV Sarsia (Celtic Sea)
RV Sarsia (E1)
RV Sarsia (E Channel Weymouth)
RV Sarsia (W Approaches)
RV Sarsia (W Channel Brest)
RV Sarsia (E5)a
RRS Discovery (NW Africa upwelling study)
RV Sarsia (W Approaches)
RV Sarsia (W Approaches)
RV Sarsia (around UK fronts Peterhead)
RV Sarsia (W Approaches)a
RV Sarsia (W Approaches)
RV Sarsia (W Approaches Douarnenez)
RV Sarsia (W Channel)a
RV Sarsia (W Channel eddies Douarnenez)
RV Sarsia (W Approaches)
RRS Challenger (Celtic Sea)
RV Oceanus (Sargasso Sea, Georges Bank)
RV Sarsia (W Approaches)a
RV Sarsia (Celtic Sea Galway)
RRS Frederick Russell (W Channel Roscoff)a
RRS Frederick Russell (Skagerrak Kristiansand)
RRS Frederick Russell (shelf edge Brest)a
RV Gyre (Gulf of Maine)
RRS Frederick Russell (Bay of Biscay)
RRS Frederick Russell (W Channel)
RRS Frederick Russell (shelf edge Penzance)a
RRS Frederick Russell (W Channel)
RRS Frederick Russell (Celtic Sea)a
RRS Frederick Russell (W Channel Penzance)
RRS Frederick Russell (shelf edge)
RRS Frederick Russell (around UK Dundee)a
RRS Charles Darwin (S North Sea, Chernobyl plume)a
RV Cirolana (NE Atlantic)
RRS Challenger (NE Atlantic and Faeroes)a
RV Argo Maine (Gulf of Maine)
RV Argo Maine (Gulf of Maine)
RV Cape Hatteras (Gulf of Maine)a
RRS Charles Darwin (NE Atlantic)a
RRS James Clark Ross (AMT2, Falklands-Grimsby)
RRS James Clark Ross (AMT6, Cape Town-Falmouth)
RRS Challenger (W Channel Plymouth)
RRS James Clark Ross (Celtic Sea)
RRS James Clark Ross (AMT 14, Falklands-Grimsby)a
RRS Charles Darwin (Celtic Sea Cork)
RRS Discovery (AMT 17, Glasgow-Port Elizabeth Azores)a
RRS James Clark Ross (Drake Passage, Falklands-Rothera)

Research cruises when Patrick was the Principal Scientist.

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well-mixed coastal water which occur around the coasts

of France and SW England (Pingree et al., 1975, 1976).
In the frontal region, where Chl a concentrations
reached a remarkable 100 mg m23, the dinoflagellate
Gyrodinium aureolum (now known as Karenia mikimotoi) was
the dominant species. The following summer, Robin
and Patrick returned to study the Gyrodinium bloom
occurring in the Western Approaches, using a submersible pump system which allowed continuous measurement of temperature, salinity, Chl a, silicate and nitrate
in surface and subsurface waters (Pingree et al., 1977).
The use of infrared satellite images alongside in situ data
provided the first confirmation that eddy structures seen
on satellite pictures of the Ushant front were intrusions
of cold, mixed water into warmer, more stratified water.
In addition, the high Gyrodinium (Karenia) counts in the
cold water patch provided evidence for upward displacement of thermocline waters within the eddy (Pingree
et al., 1979). The collaboration with Peter Baylis
(University of Dundee), who supplied the very high resolution radiometer images from the NOAA 5 satellite,
provided the first recognition of the power of having relevant satellite images while at sea. From then on satellite images, which were printed out on an old British
War Department facsimile machine, were posted to
France where the RV Sarsia docked mid-cruise to pick
them up. This was a convenient excuse for a run
ashore and the resulting activities have become the
stuff of Plymouth folklore!
In April 1979, Patrick returned to the Celtic Sea in
spring to join a cruise on RRS Challenger organized by
Mike Fasham and Phil Pugh from IOS, Wormley.
Regional surveys during the period of the spring bloom
formed the basis for developing and validating a model
of phytoplankton growth (Fasham et al., 1983). The
carbon-to-chlorophyll ratio and photo-acclimation of
the phytoplankton and grazing pressure by zooplankton
were all identified as key issues for improving our
understanding of the dynamics of natural phytoplankton populations.
Robin Pingree and Patrick had given talks on their
work in 1978 on tidal fronts at a NATO-sponsored
meeting in Sicilythis was Patricks first overseas conference. Here, he met Charlie Yentsch, who invited
Patrick to visit the Bigelow Laboratory for Ocean
Sciences, ME, USA, the following summer. At Bigelow,
Patrick formed his first links with scientists in the USA
working on phytoplankton, including Richard (Dick)
Dugdale, Ian Morris, Clarice Yentsch, Chris Garside
and Barney Balch. He participated in a cruise on the
RV Oceanus in August 1979 (Table I) to study phytoplankton growth and nitrogen dynamics in the Gulf of
Maine and Sargasso Sea. The ammonium and nitrite

profiles provided evidence for nitrification occurring at

the base of the pycnocline, below the chlorophyll
maximum. The distribution of the phytoplankton taxonomic composition suggested that active growth and
aggregation of particular motile species were as important in the formation and maintenance of the subsurface
chlorophyll maximum as passive mechanisms such as
physical mixing and the sinking properties of cells
(Holligan et al., 1984a).
Back in the UK, Patrick undertook a combined satellite and shipboard study of the Celtic and Armorican
Shelf edge, where every year since the Coastal Zone
Color Scanner (CZCS) was launched, extensive patches
of water giving strong reflectance of visible light had
been observed. Patrick and colleagues confirmed that
the high reflectance was caused by a surface bloom of
the coccolithophore Emiliania huxleyi at densities of up to
8500 cells mL21 (Holligan et al., 1983a). The reflective
water was visually turbid, with a pale turquoise colour
apparent even in aerial photographs from altitudes up
to 10 km. The fact that these blooms had not been
recognized in an area relatively well studied by oceanographers for nearly 100 years emphasized the important contribution satellite imagery could make to
investigations of large scale biological processes
(Holligan et al., 1983a).
Accurate visualization of Gyrodinium (Karenia) blooms
in satellite images required correction for atmospheric
conditions that interfered with the water-leaving
signals. The application of full atmospheric corrections
to CZCS data for the English Channel was carried out
by Michel Viollier at the University of Lille, France, as
described by Holligan et al. (Holligan et al., 1983b).
This study, to calibrate and validate ocean colour
sensors, was also Patricks first collaboration with Jim
Aiken, who oversaw the deployment of the undulating
oceanographic recorder along transects between
Plymouth and Roscoff. The drive to develop further
the atmospheric corrections led Patrick to collaborate
with Gary Hunt at Imperial College, London. Steve
Groom was appointed to a PhD studentship on the
application of procedures to estimate chlorophyll and
coccolithophore abundance (Holligan and Groom,
1986; Groom and Holligan, 1987) and this culminated
in the publication of a satellite colour atlas for the
North Sea, in which CZCS images revealed the
complex seasonal changes in optical and biological
properties of surface waters not previously resolved
through direct observations from ships (Holligan et al.,
1989). The image on the front cover of the atlas
became an icon of satellite oceanography and has only
recently been superseded as such by various SeaWiFs
images.

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In July/August 1981, Patrick led a cruise on the RRS

Frederick Russell (Table I) to compare and contrast the
plankton community composition and activity in mixed,
frontal and stratified waters of the English Channel.
Collaboration with Roger Harris, Peter J. leB. Williams,
Duncan Purdie, Richard Newell, Mike Lucas and
Annie Lindley enabled standing stocks of bacterial, phytoplankton, protozoan, micro- and meso-zooplankton
carbon to be estimated independently, alongside
measurements of photosynthesis, respiration and zooplankton ammonium excretion. Major differences were
found across the frontal boundary in the abundance
and species composition of the phytoplankton, and in
the relative importance of the different groups of heterotrophs. Holligan et al. (Holligan et al., 1984b, c) discussed the ecological implications of these biomass and
activity distribution patterns and emphasized the need
for quantitative observations on the transfer of carbon
within the different food webs that develop as a result of
differences in tidal mixing.
During the mid-1980s, through collaboration with
Peter Liss, Suzanne Turner and Gill Malin, Patrick
investigated variations in dimethylsulphide (DMS) concentrations in relation to hydrographic structures and
the distribution of phytoplankton species on the northwest European continental shelf and in the NE Atlantic.
Much of the variability in surface concentrations of
DMS and its precursor DMSP could be ascribed to the
species composition of the phytoplankton. Phaeocystis
and coccolithophores were important sources of DMS
(Holligan et al., 1987; Turner et al., 1988; Malin et al.,
1993).
In 1986, Patrick attended a meeting in Paris organised by the International Council for Science (ICSU)
and chaired by Jim McCarthy from Harvard University
to discuss interactive processes at the interface between
the atmosphere and the oceans. This initiative was one
of several that recognised the need for internationally
co-ordinated global-scale biogeochemistry programmes
to study the phenomenon of global change, and which
eventually formed the basis for the IGBP (www.igbp.
net). The implementation of IGBP began in the early
1990s with the official launch of five projects, including
the Joint Global Ocean Flux Study (JGOFS).
In 1988, Patrick began a 2-year secondment to the
Bigelow Laboratory for Ocean Sciences, ME, USA as
the Acting Director, where the 50 or so staff now
included John Cullen, Bob Guillard, and Maureen
Keller. Patrick collaborated with Steve Ackleson to
develop the use of remote sensing to detect coccolithophore blooms in the Gulf of Maine (Ackleson and
Holligan, 1989; Ackleson et al., 1994) and Barney Balch
and David Townsend to investigate the ecological and

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optical properties of these blooms (Balch et al., 1991,

1992; Townsend et al., 1994a, b; Table I). Patrick was
succeeded as Director by Denis Taylor in 1989. While
abroad, a large part of the MBA and the UK Natural
Environment Research Councils (NERC), Institute for
Marine Environmental Research (IMER), were merged
in a rationalization of NERC-funded marine institutes,
to become the Plymouth Marine Laboratory (PML).
Before his return to this changed landscape, Patrick had
successfully defended a promotion application to an
individual merit (Band 3) position in PML.
A few months after returning to PML, Patrick
attended an IGBP-sponsored meeting in Tokyo on
coastal processes, which led eventually to him becoming
Chair of the Scientific Steering Committee for the
IGBP Land-Ocean Interactions in the Coastal Zone
(LOICZ) project. He steered the nominally 10 year programme through its planning and early implementation
stages, with the International Project Office established
at the Netherlands Institute for Sea Research (NIOZ).
During the 5 year tenure of his Chairmanship, Patrick
made more than 65 international visitsequivalent to
around one trip per month. In 1995, he handed over
the LOICZ Chairmanship to Ed Gomez from the
Philippines.
Meanwhile in the UK, NERC had funded one of the
early scientific
community
programmesthe
Biogeochemical Ocean Flux Study (BOFS) as a contribution to the IGBP-sponsored JGOFS programme.
Patrick contributed a proposal for a North Atlantic
research cruise focussing on the biogeochemical consequences of blooms of the coccolithophore Emiliania
huxleyi. An extensive (250 000 km2) bloom of E. huxleyi
south of Iceland was studied in June 1991 during a
period of fine weather that allowed excellent visualization from both aircraft and satellites of its distribution
(Holligan et al., 1993a). Coccolith production (1  106
tonnes calcite-C) had a significant impact on the CO2
system, causing relative increases of up to 50 matm in
surface pCO2 (Robertson et al., 1994). This cruise
involved collaboration with Emilio Fernandez, a Marie
Curie Fellow from the University of Vigo, whose
measurements of photosynthesis and calcification
showed that significant amounts of carbon were incorporated into coccoliths in darkness and that on some
occasions, calcification in subsurface populations saturated at higher irradiances than in surface populations
(Fernandez et al., 1993).
In 1995, Patrick moved to the University of
Southampton, UK, as Professor of Oceanography. In
1998, he became the first Head of the SOES, initiating
the coming together and strengthening of the former
Departments of Oceanography and Geology. Steering

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any change process of this type is inherently demanding, but Patrick led with warmth, good humour and
clarity of vision. During his 4-year tenure as Head of
School, Patrick initiated the development of marine
molecular biology at Southampton and oversaw the
process whereby SOES became the main UK centre
for marine geophysics, as well as expanding capabilities
in marine geochemistry. Patricks legacy as a scientific
leader continues to be warmly regarded as having been
crucially instrumental in developing oceanography and
earth science at Southampton.
In his 12 years at Southampton, Patrick supervised
16 PhD students (Table II) and mentored a number of
postdoctoral staff including Toby Tyrrell, Cathy Lucas
and Emilio Maranon. The effects of coccolith light scattering on the optical properties of seawater were investigated by Toby Tyrrell. The Monte Carlo modelling
study showed that high coccolith concentrations caused
increased brightness and heating of the surface water,
increased albedo of the ocean, stronger stratification,
net cooling of the water column and decreased total
water column productivity (Tyrrell et al., 1999). At the
same time, a new interest in intertidal benthic microalgae had led to funding of a project within the
EU-funded programme ECOFLAT (Eco-Metabolism of
a Tidal Flat). Studies of algal pigment distribution on
the Molenplaat tidal flat within the Westerschelde
estuary (SW Netherlands) showed that microalgal
biomass was redistributed both horizontally and vertically by a combination of biological and physical processes (Lucas and Holligan, 1999). Chl a distribution
was highly correlated with sedimentological parameters,
with algal biomass greatest at siltier sites. Field measurements and annular flume experiments also showed that

benthic abundance and resuspension of large planktonic diatoms, such as Coscinodiscus spp., increased in
response to increases in current velocity (Lucas et al.,
2001).
In 1994, together with Jim Aiken at the Plymouth
Marine Laboratory, Patrick had written one of the two
NERC standard grants within the PRIME (Plankton
Reactivity in the Marine Environment) thematic programme which would become the AMT programme.
Patrick participated in the second AMT cruise between
the Falkland Islands and the UK, crossing the equator
on board ship for the first time in his career. His
research on oceanic phytoplankton flourished through
the opportunities afforded by the AMT. Maranon and
Holligan (Maranon and Holligan, 1999) and Maranon
et al. (Maranon et al., 2000) examined the latitudinal
and vertical distributions of phytoplankton size structure
and productivity in relation to physical, chemical and
biological variables along the AMT track between 508N
and 508S. A surprisingly large degree of variability
occurred in the Atlantic subtropical gyres, where estimated rates of primary production and phytoplankton
growth varied by as much as 8-fold, while Chl a concentrations remained relatively constant. This has major
implications for satellite-based models of global productivity, because the observed variability in biomassspecific photosynthesis was at least three times higher
than previously assumed. These studies also found a
significant negative correlation between the latitudinal
changes in the maximum Chl a normalized rate of
photosynthesis (P Bm) and those in the depth of the nitracline, emphasizing the need to include nutrient driven
changes of phytoplankton photophysiology in models of
primary productivity.

Table II: PhD students supervised by Patrick

Stephen Groom
Graham Dixon
Lesley Dowling
Isabelle Dias
Leonie Dranseld
Matthew Pinkerton
William Spooner
David Suggett
Patrick Friend
Mark Moore
Alex Poulton
Katherine Woods
Ibrahim Al-Tisan
Elena San Martin
Anna Hickman
Jackie Tweddle
Mohammed Qurban
Young-Nam Kim
Jane Heywood
Tim Adey

Imperial College (with G. Hunt)

University of Swansea (with P. Syrrett)
University of Cambridge (with N. McCave)
University of Bristol (with G. Eglinton)
University of Southampton
University of Southampton (with J. Aiken, PML)
University of Southampton
University of Southampton
University of Southampton (with M. Collins)
University of Southampton (with J. Sharples)
University of Southampton
University of Southampton (with S. Groom, PML)
University of Southampton
University of Southampton (with R. Harris, PML)
University of Southampton (with J. Sharples)
University of Southampton (with J. Sharples)
University of Southampton (with D. Hydes)
University of Southampton
University of Southampton (with M. Zubkov)
University of Southampton

99

1983
1983 1986
1985 1989
1986 1990
1995 1999
1996 1999
1996 2000
1997 2001
1997 2000
1998 2002
1998 2002
1999 2003
2001 2006
2002 2005
2002 2007
2002 2007
2002
2002
2003
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Data from four AMT cruises were used to characterize the patterns of phytoplankton size structure and productivity in temperate, oligotrophic, upwelling and
equatorial regions (Maranon et al., 2001). On average,
picophytoplankton (0.2 2 mm) accounted for 56 and
71% of the total integrated carbon fixation and autotrophic biomass, respectively, and most of the latitudinal
variability in total photoautotrophic biomass and production was driven by changes in this size class. In temperate and coastal upwelling regions, small (,2 mm)
and large (.2 mm) phytoplankton accounted for a proportion of total biomass that was similar to their proportion of total productivity. In contrast, in the
oligotrophic and equatorial regions, large phytoplankton tended to account for a fraction of the total primary
production that was significantly higher than their proportion of total biomass. Interestingly, in the equatorial
upwelling, the increased biomass and production was
not associated with a corresponding change in the phytoplankton size structure (Maranon et al., 2001).
Other related work on phytoplankton pigment distributions and absorption properties was continued in collaboration with Ray Barlow and Stuart Gibb during
AMT (Gibb et al., 2000; Barlow et al., 2002, 2004) and
through supervision of PhD students Mark Moore and
David Suggett (Table II) who investigated the variability
and environmental controls on phytoplankton physiology using fast-repetition-rate-fluorometry (Moore
et al., 2003, 2006; Suggett et al., 2001, 2006).
In 2000, the first overview paper of the AMT programme was published (Aiken et al., 2000) and Patrick
became one of the main co-investigators of further
funding for AMT, through the submission of a NERC
marine/atmospheric consortium proposal. This subsequently funded 45 researchers (including 9 students) in
6 UK research establishments to undertake six 40 day,
13 500 km transect cruises between 2003 and 2005
(Robinson et al., 2006). Patrick was the Principal Scientist
on two of these cruises (Fig. 1; Table I) and at the time of
going to press, the AMT programme has enabled collaboration with 27 research groups from 10 countries and
resulted in 160 publications (www.amt-uk.org).
Patricks research within the NERC Consortium
phase of the AMT programme included the study of
coccolithophore distribution and activity, the impact of
coccolithophores on oceanic carbonate budgets and the
relative magnitude of phytoplankton photosynthesis, calcification and silicification. Simultaneous measurements
of the biogenic production of calcite (calcification), opal
(silicification) and organic carbon ( photosynthesis) in
the upper ocean between 498N and 428S in the
Atlantic Ocean, confirmed the light dependency of calcification and photosynthesis, and the substrate

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Fig. 1. Oceanographic contemplation. Photo courtesy of David

Suggett.

dependency of silicification (Poulton et al., 2006). Rates

of calcification were generally twice those of silicification, whereas rates of organic carbon production were
at least an order of magnitude higher than the rates of
calcification. The average turnover times for calcite and
phytoplankton carbon were 3 days, whereas average
turnover times for opal and particulate organic carbon
were 10 days (Poulton et al., 2006). The comparison of
surface production rates with sediment trap data confirmed that 70% of the calcite is dissolved in the
upper 2 3 km, and only a tiny proportion of total
organic carbon (,2%) reaches the deep ocean (Poulton
et al., 2006). By comparing these measurements with
similar data from other oceanic settings, Poulton et al.
(Poulton et al., 2007) were able to show the considerable
regional variability in the positive correlations between
oceanic calcification and photosynthesis, Chl a or
calcite concentration, emphasizing that both cocolithophore physiology and community ecology dictate the
biogeochemical impact of coccolithophores.
During this time, Patrick also renewed his interests in
shelf seas by undertaking three cruises between 1999
and 2005 (Table I) with Jonathan Sharples and John
Simpson to investigate the role of tidal mixing on phytoplankton growth by supplying bottom-layer nutrients
into the subsurface chlorophyll maximum. An average

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nitrate flux into the thermocline generated by tidal

mixing was estimated to be capable of supporting new
phytoplankton growth at a rate of 160 mg C m22
day21 (Sharples et al., 2001). This has important implications for attempts to quantify primary production in
shelf seas as neither remote sensing of the colour of the
sea surface nor standard incubation techniques would
be able to account for the variability in production rates
at the subsurface maximum caused by this tidal
pumping. In a first assessment of this variability in
nitrate supply associated with mixing changes during
the spring-neap tidal cycle at the Celtic Sea shelf break,
Sharples et al. (Sharples et al., 2007) measured a neap
tide daily mean nitrate flux of 1.3 mmol m22 day21
and estimated the spring tide nitrate flux to be as high
as 9 mmol m22 day21.
In 2005, Patrick retired from the University of
Southampton. Since 1970 he has published 91 peer
reviewed manuscripts, including 31 first author papers
and 5 articles in Nature, with a total of 4012 citations
and an average citation rate per publication of 45 (Web
of Science, 27 November 2007; Fig. 2). Following his
retirement, Patrick became the President of the Council
of the SAHFOSthe international charity which operates the Continuous Plankton Recorder (CPR) Survey.

In 2006, Patrick was awarded the Plymouth Marine

Sciences Silver Medal. This prestigious award was
established in 1986 and recognizes and rewards outstanding contributions to marine science. It was particularly fitting since much of Patricks most significant
work had been conducted when he was based in
Plymouth. In his Plymouth Medal Lecture, Patrick
stressed with conviction his belief in the importance of
observations, while cautioning against a reliance on
models alone: an elegant synthesis of much of his lifes
work. In this spirit he undertook his last research cruise,
at the end of 2006, to investigate the distributions of calcareous and siliceous phytoplankton across the Drake
passage in the Southern Ocean.
In trying to summarize Patricks scientific career into
a very few pages, we are conscious of the superficial
attention we have given to every aspect of his professional life. We are also conscious that Patricks other
colleagues would doubtless choose to emphasize different aspects of his science than we have done: this is the
inevitable problem of the biographer. Nevertheless, we
hope we have captured the milestones and important
landmarks of Patricks career. In initiating this special
issue of the Journal of Plankton Research, we hope we have
also given the opportunity to some of Patricks colleagues to review and celebrate his many scientific collaborations. Special thanks are due to the many referees
who helped to process these papers, often working to
very tight deadlines, to Roger Harris for advice, Aditee
Mitra for handling the papers and to Kevin Flynn for
editing them.
Finally, we would like to record our own personal
thanks to Patrick for the professional and personal interactions we have enjoyed, and on behalf of all those who
know Patrick, we wish him continued success, good
fortune and smooth seas.

P U B L I C AT I O N S O F PAT R I C K
M. HOLLIGAN (IN
C H RO N O LO G I C A L O R D E R )

Fig. 2. Publications and citations per year.

Holligan, P. M. (1971) Routine analysis by gas liquid

chromatography of soluble carbohydrates in extracts
of plant tissues. I. A review of techniques used for
the separation, identification and estimation of carbohydrates by gas liquid chromatography. New Phytol.,
70, 239 269.
Holligan, P. M. and Drew, E. A. (1971) Routine
analysis by gas liquid chromatography of soluble
carbohydrates in extracts of plant tissues. II.
Quantitative analysis of standard carbohydrates and

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the separation and estimation of soluble sugars and

polyols from a variety of plant tissues. New Phytol., 70,
271 297.
Holligan, P. M. and Jennings, D. H. (1972)
Carbohydrate metabolism in the fungus Dendryphiella
salina. I. Changes in the levels of soluble carbohydrates during growth. New Phytol., 71, 569 582.
Holligan, P. M. and Jennings, D. H. (1972)
Carbohydrate metabolism in the fungus Dendryphiella
salina. II. The influence of different carbon and nitrogen sources on the accumulation of mannitol and
arabitol. New Phytol., 71, 583 594.
Holligan, P. M. and Jennings, D. H. (1972)
Carbohydrate metabolism in the fungus Dendryphiella
salina. III. The effect of the nitrogen sources on the
metabolism of [1 14C] and [6 14C] glucose. New
Phytol., 71, 1119 1133.
Holligan, P. M. and Jennings, D. H. (1972)
Carbohydrate metabolism in the fungus Dendryphiella
salina. IV. Acetate assimilation. New Phytol., 72, 315
319.
Holligan, P. M. and Jennings, D. H. (1972)
Unexpected labelling patterns from radioactive
sugars fed to plants containing mannitol.
Phytochemistry, 11, 3447 3451.
Holligan, P. M. and Lewis, D. H. (1973) The soluble
carbohydrates of Aspergillus clavatus. J. Gen. Microbiol.,
75, 155 159.
McGee, E. E. M., Holligan, P. M., Fung, A. K. and
Lewis, D. H. (1973) Maintenance of the rust, Puccinia
poarum, on its alternate hosts under controlled conditions. New Phytol., 72, 937 945.
Holligan, P. M., Chen, C. and Lewis, D. H. (1973)
Changes in the carbohydrate composition of leaves
of Tussilago farfara during infection by Puccinia poarum.
New Phytol., 72, 947 955.
Holligan, P. M., McGee, E. E. M. and Lewis, D. H.
(1974) Quantitative determination of starch and
glycogen and their metabolism in leaves of Tussilago
farfara during infection by Puccinia poarum. New Phytol.,
73, 873 879.
Holligan, P. M., Chen, C., McGee, E. E. M. and
Lewis, D. H. (1974) Carbohydrate metabolism in
healthy and rusted leaves of coltsfoot. New Phytol., 73,
881 888.
Gilbert, O. L., Holligan, P. M. and Holligan,
M. S. (1974) The flora of North Rona 1972. Trans.
Bot. Soc. Edinb., 42, 43 68.
Holligan, P. M. and Gooday, G. W. (1975) Symbiosis
in Convoluta roscoffensis. Symp. Soc. Exp. Biol., 29, 205
227.
Pingree, R. D., Pugh, P. R., Holligan, P. M. and
Forster, G. R. (1975) Summer phytoplankton blooms

30

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and red tides along tidal fronts in the approaches to

the English Channel. Nature, Lond., 258, 672 677.
Pingree, R. D., Holligan, P. M., Mardell, G. T. and
Head, R. N. (1976) The influence of physical stability
on spring, summer and autumn phytoplankton
blooms in the Celtic Sea. J. Mar. Biol. Assoc., 56,
845 873.
Pingree, R. D., Holligan, P. M. and Head,
R. N. (1977) The survival of dinoflagellate blooms in
the western English Channel. Nature, Lond., 265,
266 269.
Holligan, P. M. and Harbour, D. S. (1977). The vertical distribution and succession of phytoplankton in
the western English Channel in 1975 and 1976.
J. Mar. Biol. Assoc., 57, 1075 1093.
Pingree, R. D., Holligan, P. M. and Mardell, G. T.
(1978) The effects of vertical stability on phytoplankton distributions in the summer on the northwest
European Shelf. Deep-Sea Res., 25, 1011 1028.
Holligan, P. M., Pingree, R. D., Pugh, P. R. and
Mardell, G. T. (1978) The hydrography and plankton
of the eastern English Channel in March 1976. Ann.
Biol., 33, 69 71.
Mackie, P. R., Hardy, R., Butler, E. I., Holligan,
P. M. and Spooner, M. F. (1978). Early samples of oil
in water and some analyses of zooplankton. Mar.
Pollut. Bull., 9, 296 297.
Pingree, R. D., Holligan, P. M. and Mardell, G. T.
(1979) Phytoplankton growth and cyclonic eddies.
Nature, Lond., 278, 245 247.
Gilbert, O. L. and Holligan, P. M. (1979) Puccinellia capillaris (Liljebl.) Jans  P. maritima (Huds.) Parl. on North
Rona, Outer Hebrides. Watsonia, 12, 338339.
Holligan, P. M., Maddock, L. and Dodge, J. D.
(1980) The distribution of dinoflagellates around the
British Isles in July, 1977: a multivariate analysis.
J. Mar. Biol. Assoc., 60, 851 867.
Holligan, P. M. (1981) Biological implications of
fronts on the northwest European continental shelf.
Philos. Trans. R. Soc. Lond., A 302, 547 562.
Pingree, R. D., Mardell, G. T., Holligan, P. M.,
Griffiths, D. K. and Smithers, J. (1982) Celtic Sea
and Armorican current structure and the vertical distributions of temperature and chlorophyll. Contin.
Shelf. Res., 1, 99 116.
Pingree, R. D., Holligan, P. M., Mardell, G. T. and
Harris, R. P. (1982) Vertical distributions of plankton
in the Skagerrak in relation to doming of the seasonal thermocline. Contin. Shelf. Res., 1, 209 219.
Fasham, M. J. R., Holligan, P. M. and Pugh,
P. R. (1983). The spatial and temporal development
of the spring phytoplankton bloom in the Celtic Sea,
April 1979. Prog. Oceanogr., 12, 87 145.

C. ROBINSON AND N. J. P. OWENS

PATRICK MICHAEL HOLLIGAN: A SHORT BIOGRAPHY

Holligan, P. M., Viollier, M., Harbour, D. S.,

Camus, P. and Champagne-Phillippe, M. (1983a)
Satellite and ship studies of coccolithophore production along a continental shelf edge. Nature, Lond.,
304, 339 342.
Holligan, P. M., Viollier, M., Dupuoy, C. and Aiken,
J. (1983b) Satellite studies on the distributions of
chlorophyll and dinoflagellate blooms in the western
English Channel. Contin. Shelf. Res., 2, 81 96.
Holligan, P. M., Balch, W. M. and Yentsch,
C. M. (1984a) The significance of subsurface chlorophyll, nitrite and ammonium maxima in relation
to nitrogen for phytoplankton growth in stratified
waters of the Gulf of Maine. J. Mar. Res., 42, 1051
1073.
Holligan, P. M., Harris, R. P., Newell, R. C.,
Harbour, D. S., Head, R. N., Linley, E. A. S.,
Lucas, M. I., Tranter, P. G. R. and Weekley,
C. M. (1984b) Vertical distribution and partitioning
or organic carbon in mixed, frontal and stratified
waters of the English Channel. Mar. Ecol. Prog. Ser.,
14, 111 127.
Holligan, P. M., Williams, P. J., Le, B., Purdie, D. and
Harris, R. P. (1984c) Photosynthesis, respiration and
nitrogen supply of plankton populations in stratified,
frontal and tidally mixed waters. Mar. Ecol. Prog. Ser.,
17, 201 213.
McCarthy, P. M and Holligan, P. M. (1984)
Inishvickillane, Co. Kerry: an important lichen site.
Ir. Nat. J., 21, 266 274.
Holligan, P. M., Pingree, R. D. and Mardell,
G. T. (1985). Oceanic solitons, nutrient pulses
and phytoplankton growth. Nature, Lond., 314, 348350.
Holligan, P. M. and Groom, S. B. (1986).
Phytoplankton distributions along the shelf break.
Proc. R. Soc. Edinb., 88B, 239 263.
Holligan, P. M., Turner, S. M. and Liss, P. S. (1987)
Measurements of dimethyl sulphide in frontal
regions. Contin. Shelf. Res., 7, 213 224.
Holligan, P. M. (1987) The physical environment of
phytoplankton blooms in the N.E. Atlantic. Rapp. P.-v.
Cons. Perm. Int. Explor. Mer., 187, 9 18.
Groom, S. B. and Holligan, P. M. (1987) Remote
sensing of coccolithophore blooms. Adv. Space Res.,
7(2), 73 78.
Turner, S. M., Malin, G., Liss, P. S., Harbour, D. S.
and Holligan, P. M. (1988) The seasonal variation of
dimethyl sulphide and dimethylsulphoniopropionate
concentrations in nearshore waters. Limnol. Oceanogr.,
33, 364 375.
Ackleson, S., Balch, W. M. and Holligan, P. M.
(1988) White waters of the Gulf of Maine.
Oceanography, 1(2), 18 22.

Holligan, P. M. (1989) Primary productivity in the

shelf seas of northwest Europe. Adv. Bot. Res., 16,
193 252.
Dixon, G. K. and Holligan, P. M. (1989) Studies on
the growth and nitrogen assimilation of the bloom
dinoflagellate Gyrodinium aureolum Hulbert. J. Plankton
Res., 11, 105 118.
Ackleson, S. and Holligan, P. M. (1989) AVHRR
observations of a Gulf of Maine coccolithophore
bloom. Photogram. Eng. Rem. Sens., 55, 473 474.
Holligan, P. M., Aarup, T. and Groom, S. (1989) The
North Sea: satellite colour atlas. Contin. Shelf Res., 9,
667 765.
Aarup, T., Groom, S. and Holligan, P. M. (1989)
CZCS imagery of the North Sea. Adv. Space Res., 9,
443 451.
Aarup, T., Groom, S. and Holligan, P. M. (1990) The
processing and interpretation of North Sea CZCS
imagery. Neth. J. Sea Res., 25, 3 9.
Williamson, P. and Holligan, P. M. (1990) Ocean productivity and climate change. Trends. Ecol. Evol., 5,
299 303.
Balch, W. M., Holligan, P. M., Ackleson, S. G. and
Voss, K. J. (1991) Biological and optical properties of
mesoscale coccolithophore blooms in the Gulf of
Maine. Limnol. Oceanogr., 36, 629 643.
Holligan, P. M. and Reiners, W. A. (1992) Predicting
the responses of the coastal zone to global change.
Adv. Ecol. Res., 22, 211 255.
Balch, W. M., Holligan, P. M. and Kilpatrick, K. A.
(1992) Calcification, photosynthesis and growth of
the bloom-forming coccolithophore, Emiliania huxleyi.
Contin. Shelf Res., 12, 1353 1374.
Aiken, J., Moore, G. and Holligan, P. M. (1992)
Remote sensing of oceanic biology in relation to
global climate change. J. Phycol., 28, 570 590.
Malin, G., Turner, S., Liss, P., Holligan, P. and
Harbour, D. (1993) Dimethyl sulphide and dimethylsulphonioproprionate in the north east Atlantic
during the summer coccolithophore bloom. Deep-Sea
Res. I, 40, 1487 1508.
Fernandez, E., Boyd, P., Holligan, P. M. and
Harbour, D. S. (1993) Production of organic and
inorganic carbon within a large scale coccolithophore bloom in the North-east Atlantic Ocean. Mar.
Ecol. Prog. Ser., 97, 271 285.
Balch, W. M., Kilpatrick. K., Holligan, P. M. and
Cucci, T. L. (1993) Coccolith production and detachment by Emiliania huxleyi (Prymnesiophyceae).
J. Phycol., 29, 566 575.
Holligan, P. M., Fernandez, E., Aiken, J.,
Balch, W. M., Boyd, P., Burkill, P. H., Finch, M.,
Groom, S. B., Malin, G., Muller, K., Purdie, D. A.,

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Robinson, C., Trees, C. C., Turner, S. M. and van

der Wal, P. (1993a) A biogeochemical study of the
coccolithophore, Emiliania huxleyi, in the North
Atlantic. Global Biogeochem. Cycles, 7, 879 900.
Holligan, P. M., Groom, S. B. and Harbour,
D. S. (1993b) What controls the distribution of the
coccolithophore, Emiliania huxleyi, in the North Sea?
Fish. Oceanogr., 2, 175 183.
Robertson, J. E., Robinson, C., Turner, D. R.,
Holligan, P., Watson, A. J., Boyd, P., Fernandez,
E. and Finch, M. (1994) The impact of a coccolithophore bloom on oceanic carbon uptake in the northeast Atlantic during summer 1991. Deep-Sea Res. I.,
41, 297 314.
Liss, P. S., Malin, G., Turner, S. M. and
Holligan, P. M. (1994) Dimethyl sulphide and
Phaeocystis: a review. J. Mar. Syst., 5, 41 53.
Ackleson,
S.
G.,
Balch,
W.
M.
and
Holligan, P. M. (1994) Response of water-leaving
radiance to particulate calcite and chlorophyll a concentrations: a model for Gulf of Maine coccolithophore blooms. J. Geophys. Res., 99, 7483 7499.
Townsend, D. W., Keller, M. D., Holligan, P. M.,
Ackleson, S. G. and Balch, W. M. (1994) Blooms of
the coccolithophore Emiliania huxleyi with respect to
hydrography in the Gulf of Maine. Contin. Shelf Res.,
14, 979 1000.
Townsend, D. W., Cammen, L. M., Holligan, P.,
Campbell, D. E. and Pettigrew, N. R. (1994) Causes
and consequences of variability in the timing of
spring phytoplankton blooms. Deep-Sea Res. I., 41,
747 765.
Fernandez, E., Balch, W. M., Maranon, E. and
Holligan, P. M. (1994) High rates of lipid biosynthesis
in cultured, mesocosom and coastal populations of
the coccolithophore Emiliania huxleyi. Mar. Ecol. Prog.
Ser., 114, 13 22.
Hays, G. C., Warner, A. J., John, A.W. G., Harbour,
D. S. and Holligan, P. M. (1995) Coccolithophores
and the Continuous Plankton Recorder survey.
J. Mar. Biol. Assoc., 75, 503 506.
Holligan, P. M. and Robertson, J. E. (1996)
Significance of ocean carbonate budgets for the
global carbon cycle. Global Change Biol., 2, 85 95.
Balch, W. M., Kilpatrick, K. A, Holligan, P.,
Harbour, D. and Fernandez, E. (1996) The 1991 coccolithophore bloom in the central North Atlantic. 2
Relating optics to coccolith concentration. Limnol.
Oceanogr., 44, 1684 1696.
Tyrrell, T. and Holligan, P. M. (1997) How do coccoliths affect light in the sea? Proc. Soc. Photo Opt. Instrum.
Eng., 2963, 554 559.

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Tyrrell, T., Holligan, P. M. and Mobley, C. D. (1999)

Optical impacts of oceanic coccolithophore blooms.
J. Geophys. Res., 104, 3223 3241.
Maranon, E. and Holligan, P. M. (1999)
Photosynthetic parameters of phytoplankton from
50oN to 50oS in the Atlantic Ocean. Mar. Ecol. Prog.
Ser., 176, 191 203.
Lucas, C. H. and Holligan, P. M. (1999) Nature
and ecological implications of algal pigment
diversity on the Molenplaat tidal flat (Westerschelde
Estuary, SW Netherlands). Mar. Ecol. Prog. Ser., 180,
51 64.
Maranon, E., Holligan, P. M., Varela, M., Mourino,
B. and Bale, A. J. (2000) Basin-scale variability of
phytoplankton biomass, production and growth in
the Atlantic Ocean. Deep-Sea Res. I., 47, 825 857.
Gibb, S. W., Barlow, R. G., Cummings, D. G., Rees,
N. W., Trees, C. C., Holligan, P. and Suggett,
D. (2000) Surface phytoplankton pigment distributions in the Atlantic Ocean: an assessment of
basin scale variability between 508N and 508S. Prog.
Oceanog., 45, 339 368.
Aiken, J., Rees, N., Hooker, S., Holligan, P., Bale, A.,
Robins, D., Moore, G., Harris, R. and Pilgrim,
D. (2000) The Atlantic Meridional Transect: overview
and synthesis of data. Prog. Oceanog., 45, 257312.
Lucas, C. H., Banham, C. and Holligan,
P. M. (2001) Benthic-pelagic exchange of microalgae
at a tidal flat. 2. Taxonomic analysis. Mar. Ecol. Prog.
Ser., 212, 29 52.
Sharples, J., Moore, C. M., Rippeth, T. P., Holligan,
P. M., Hydes, D. J., Fisher, N. R. and Simpson,
J. H. (2001) Phytoplankton distribution and survival
in the thermocline. Limnol. Oceanogr., 46, 486 496.
Suggett, D., Kraay, G., Holligan, P., Davey, M.,
Aiken, J. and Geider, R. (2001) Assessment of photosynthesis in a spring cyanobacterial bloom by use of
a fast repetition rate fluorometer. Limnol. Oceanogr., 46,
802 810.
Maranon, E., Holligan, P. M., Barciela, R., Gonzalez,
N., Mourino, B., Pazo, M. J. and Varela, M. (2001)
Patterns of phytoplankton size structure and productivity in contrasting open-ocean environments. Mar.
Ecol. Prog. Ser., 216, 4356.
Barlow, R. G., Aiken, J., Holligan, P. M., Cummings,
D. G., Maritorena, S. and Hooker, S. (2002)
Phytoplankton pigment and absorption characteristics along meridional transects in the Atlantic
Ocean. Deep-Sea Res. 1, 49, 637 660.
Moore, C. M., Suggett, D., Holligan, P. M.,
Sharples, J., Abraham, E. R., Lucas, M. I., Rippeth,
T. P., Fisher, N. R., Simpson, J. H. and Hydes,

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PATRICK MICHAEL HOLLIGAN: A SHORT BIOGRAPHY

D. J. (2003) Physical controls on phytoplankton physiology at a shelf sea front: a fast repetition-rate fluorometer based field study. Mar. Ecol. Prog. Ser. 259,
29 45.
Friend, P. L., Collins, M. B., and Holligan,
P. M. (2003) Day night variation of intertidal
flat sediment properties in relation to sediment stability. Estuarine Coast. Shelf Sci. 58, 663 675.
Barlow, R. G., Aiken, J., Moore, G. F., Holligan,
P. M. and Lavender, S. (2004) Pigment adaptations
in surface phytoplankton along the eastern boundary
of the Atlantic Ocean. Mar. Ecol. Prog. Ser., 281,
13 26.
Chuck, A., Tyrrell, T., Totterdell, I. J. and Holligan,
P. M. (2005) The oceanic response to carbon emissions over the next century: investigation using three
ocean carbon cycle models. Tellus 57B, 70 86.
Sharples, J. and Holligan, P. M. (2006)
Interdisciplinary studies in the Celtic Sea. In
Robinson, A. R. and Brink, K. (Eds), The Sea. Vol
14B, Harvard Univ., pp. 1003 1031.
Moore, C. M., Suggett, D. J., Hickman, A. E., Kim,
Y.-M., Tweddle, J. F., Sharples, J., Geider, R. J. and
Holligan, P. M. (2006) Phytoplankton photoacclimation and photoadaptation in response to environmental gradients in a shelf sea. Limnol. Oceanogr. 51,
936 949.
Robinson, C., Poulton, A. J., Holligan, P. M., Baker,
A. R., Forster, G., Gist, N., Jickells, T. D., Malin, G.,
Upstill-Goddard, R., Williams, R. G., Woodward,
E. M. S. and Zubkov, M. V. (2006) The Atlantic
Meridional Transect (AMT) Programme: a contextual view 1995 2005. Deep-Sea Res. II., 53, 1483
1513.
Heywood, J., Zubkov, M. V., Tarran, G. A., Fuchs,
B. M. and Holligan, P. M. (2006) Prokaryoplankton
standing stocks in oligotrophic gyre and equatorial
provinces of the Atlantic Ocean: evaluation of interannual variability. Deep-Sea Res. II., 53, 1528 1545.
Suggett, D., Moore, C. M., Maranon, E., Omachi,
C., Varela, R. A., Aiken, J. and Holligan,
P. M. (2006) Photosynthetic electron turnover in the
tropical and subtropical Atlantic Ocean. Deep-Sea Res.
II., 53, 1571 1590.
Poulton, A. J., Sanders, R., Holligan, P. M.,
Stinchcombe, M. C., Adey, T. R., Brown,
L. and Chamberlain, K. (2006a) Phytoplankton mineralization in the tropical and subtropical Atlantic
Ocean. Global Biogeochem. Cycles, 20, GB4002, doi:
10.1029/2006GB002712, 2006.
Poulton, A., Holligan, P. M., Hickman, A., Kim,
Y.-N., Adey, T. R., Stinchcombe, M. C., Holeton, C.,
Root, S. and Woodward, M. S. (2006b)

Phytoplankton carbon fixation, chlorophyll biomass

and diagnostic pigments in the Atlantic Ocean.
Deep-Sea Res. II., 53, 1591 1608.
Poulton, A. J., Adey, T. R., Balch, W. M. and
Holligan, P. M. (2007) Relating coccolithophore calcification rates to phytoplankton community
dynamics: regional differences and implications for
carbon export. Deep-Sea Res. II., 54, 538 557.
Sharples, J., Tweddle, J. F., Green, J. A. M., Palmer,
M. R., Kim, Y.-N., Hickman, A. E., Holligan, P. M.,
Moore, C. M., Rippeth, T. P. and Simpson,
J. H. (2007) Spring-neap modulation of internal tide
mixing and vertical nutrient fluxes at a shelf edge in
summer. Limnol. Oceanogr., 52(5), 1735 1747.

AC K N OW L E D G E M E N T S
Many thanks to everyone who contributed information
for this biography, particularly Peter Burkill, Roger
Harris, Alan Kemp, Duncan Purdie, Andrew Roberts
and Dave Suggett. Especial thanks go to Patrick
himself, first for inspiring the conference held in his
honour, (Plymouth, April 2008) and this journal special
issue, secondly for giving up a day of his time to reminisce on his scientific career with us and thirdly for
reviewing the first draft of this biography.

REFERENCES
Ackleson, S. and Holligan, P. M. (1989) AVHRR observations of a
Gulf of Maine coccolithophore bloom. Photogram. Eng. Rem. Sens.,
55, 473 474.
Ackleson, S. G., Balch, W. M. and Holligan, P. M. (1994) Response of
water-leaving radiance to particulate calcite and chlorophyll a concentrations: a model for Gulf of Maine coccolithophore blooms.
J. Geophys. Res., 99, 7483 7499.
Aiken, J., Rees, N., Hooker, S. et al. (2000) The Atlantic Meridional
Transect: overview and synthesis of data. Prog. Oceanog., 45,
257 312.
Balch, W. M., Holligan, P. M., Ackleson, S. G. et al. (1991) Biological
and optical properties of mesoscale coccolithophore blooms in the
Gulf of Maine. Limnol. Oceanogr, 36, 629 643.
Balch, W. M., Holligan, P. M. and Kilpatrick, K. A. (1992)
Calcification, photosynthesis and growth of the bloom-forming coccolithophore Emiliania huxleyi. Contin. Shelf Res, 12, 1353 1374.
Barlow, R. G., Aiken, J., Holligan, P. M. et al. (2002) Phytoplankton
pigment and absorption characteristics along meridional transects
in the Atlantic Ocean. Deep-Sea Res. I, 49, 637660.
Barlow, R. G., Aiken, J., Moore, G. F. et al. (2004) Pigment adaptations
in surface phytoplankton along the eastern boundary of the Atlantic
Ocean. Mar. Ecol. Prog. Ser., 281, 1326.
Fasham, M. J. R., Holligan, P. M. and Pugh, P. R. (1983) The spatial
and temporal development of the spring phytoplankton bloom in
the Celtic Sea, April 1979. Prog. Oceanog., 12, 87 145.

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Fernandez, E., Boyd, P., Holligan, P. M. et al. (1993) Production of

organic and inorganic carbon within a large scale coccolithophore
bloom in the North-east Atlantic Ocean. Mar. Ecol. Prog. Ser., 97,
271285.
Gibb, S. W., Barlow, R. G., Cummings, D. G. et al. (2000) Surface
phytoplankton pigment distributions in the Atlantic Ocean: an
assessment of basin scale variability between 508N and 508S. Prog.
Oceanog., 45, 339 368.
Groom, S. B. and Holligan, P. M. (1987) Remote sensing of coccolithophore blooms. Adv. Space Res., 7, 7378.
Holligan, P. M. and Groom, S. B. (1986) Phytoplankton distributions
along the shelf break. Proc. R. Soc. Edinb., 88B, 239 263.
Holligan, P. M., Viollier, M., Harbour, D. S. et al. (1983a) Satellite and
ship studies of coccolithophore production along a continental shelf
edge. Nature, Lond., 304, 339 342.
Holligan, P. M., Viollier, M., Dupuoy, C. et al. (1983b) Satellite studies
on the distributions of chlorophyll and dinoflagellate blooms in the
western English Channel. Contin. Shelf. Res., 2, 81 96.
Holligan, P. M., Balch, W. M. and Yentsch, C. M. (1984a) The significance of subsurface chlorophyll, nitrite and ammonium maxima in
relation to nitrogen for phytoplankton growth in stratified waters of
the Gulf of Maine. J. Mar. Res., 42, 1051 1073.
Holligan, P. M., Harris, R. P., Newell, R. C. et al. (1984b) Vertical distribution and partitioning or organic carbon in mixed, frontal and
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Holligan, P. M., Williams, P. J., Le, B. et al. (1984c) Photosynthesis, respiration and nitrogen supply of plankton populations in stratified,
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Holligan, P. M., Turner, S. M. and Liss, P. S. (1987) Measurements of
dimethyl sulphide in frontal regions. Contin. Shelf. Res., 7, 213 224.

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Moore, C. M., Suggett, D., Holligan, P. M. et al. (2003) Physical controls

on phytoplankton physiology at a shelf sea front: a fast repetition-rate
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the approaches to the English Channel Nature, Lond. 258, 672677.
Pingree, R. D., Holligan, P. M., Mardell, G. T. and Head, R. N.
(1976) The influence of physical stability on spring, summer and
autumn phytoplankton blooms in the Celtic Sea. J. Mar. Biol. Assoc.,
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Pingree, R. D., Holligan, P. M. and Mardell, G. T. (1979)
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Poulton, A. J., Sanders, R., Holligan, P. M. et al. (2006) Phytoplankton
mineralization in the tropical and subtropical Atlantic Ocean. Global
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of a coccolithophore bloom on oceanic carbon uptake in the northeast Atlantic during summer 1991. Deep-Sea Res. I., 41, 297 314.

Holligan, P. M., Aarup, T. and Groom, S. (1989) The North Sea: satellite colour atlas. Contin. Shelf Res., 9, 667 765.

Robinson, C., Poulton, A. J., Holligan, P. M. et al. (2006) The Atlantic

Meridional Transect (AMT) Programme: a contextual view 1995
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Holligan, P. M., Fernandez, E., Aiken, J. et al. (1993a) A biogeochemical study of the coccolithophore, Emiliania huxleyi, in the North
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Sharples, J., Moore, C. M., Rippeth, T. P. et al. (2001) Phytoplankton

distribution and survival in the thermocline. Limnol. Oceanogr., 46,
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Holligan, P. M., Groom, S. B. and Harbour, D. S. (1993b) What controls the distribution of the coccolithophore, Emiliania huxleyi, in the
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Sharples, J., Tweddle, J. F., Green, J. A. M. et al. (2007) Spring-neap

modulation of internal tide mixing and vertical nutrient fluxes at a
shelf edge in summer. Limnol. Oceanogr., 52, 1735 1747.

Lucas, C. H. and Holligan, P. M. (1999) Nature and ecological implications

of algal pigment diversity on the Molenplaat tidal flat (Westerschelde
Estuary, SW Netherlands). Mar. Ecol. Prog. Ser., 180, 5164.

Suggett, D., Kraay, G., Holligan, P. et al. (2001) Assessment of photosynthesis in a spring cyanobacterial bloom by use of a fast repetition
rate fluorometer. Limnol. Oceanogr., 46, 802 810.

Lucas, C. H., Banham, C. and Holligan, P. M. (2001) Benthic-pelagic

exchange of microalgae at a tidal flat. 2. Taxonomic analysis. Mar.
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Suggett, D., Moore, C. M., Maranon, E. et al. (2006) Photosynthetic

electron turnover in the tropical and subtropical Atlantic Ocean.
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Malin, G., Turner, S., Liss, P. et al. (1993) Dimethyl sulphide and
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summer coccolithophore bloom. Deep-Sea Res. I., 40, 14871508.

Townsend, D. W., Cammen, L. M., Holligan, P. et al. (1994a) Causes

and consequences of variability in the timing of spring phytoplankton blooms. Deep-Sea Res. I., 41, 747 765.

Maranon, E. and Holligan, P. M. (1999) Photosynthetic parameters of

phytoplankton from 508N to 508S in the Atlantic Ocean. Mar. Ecol.
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Townsend, D. W., Keller, M. D., Holligan, P. M. et al. (1994b) Blooms

of the coccolithophore Emiliania huxleyi with respect to hydrography
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Maranon, E., Holligan, P. M., Varela, M. et al. (2000) Basin-scale

variability of phytoplankton biomass, production and growth in the
Atlantic Ocean. Deep-Sea Res. I., 47, 825 857.

Turner, S. M., Malin, G., Liss, P. S. et al. (1988) The seasonal variation
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Maranon, E., Holligan, P. M., Barciela, R. et al. (2001) Patterns of

phytoplankton size structure and productivity in contrasting openocean environments. Mar. Ecol. Prog. Ser., 216, 43 56.

Tyrrell, T., Holligan, P. M. and Mobley, C. D. (1999) Optical impacts

of oceanic coccolithophore blooms. J. Geophys. Res., 104,
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