384 John Woods
ples, plankton species that falI into the mindless category are responsible for toxic
algal blooms, eutrophication, cholera epidemics, and the biologic al pump in the
global carbon cycle (and therefore global warming in the 21 st century and at the
end ofice ages). Primitive equation modelling is the most promising technique for
predicting these phenomena.
18.4.4 Reproducible laboratory experiments
It will first be necessary to devise reproducible laboratory experiments to establish biological primitive equations for each of the basic functions of the physiology
and behaviour of plankton. Because these organisms are tiny, the experiments will
most likely be performed on populations rather than individuals. However, that is
not a problem ifthe individuals in the populations are identical, being clones from
a single egg, and therefore identical geneticalIy. Dr.Carlotti has drawn my attention
to the existence of significant intra-population variation in experiments with plankton from the same clutch of eggs, rather than from the same egg.
The BPEs extracted from such experiments describe aspects of the physiology
and behaviour of a single organism acting in response to its ambient environment
(which will be carefulIy controlled in the experiment). Note that the ambient environment includes physical, chemical and biological variables. The ambient biological environment experienced by a single plankter comprises the local
concentration and biologic al state of its prey, its predators, its own species and
competitors. Experiments designed to produce biological primitive equations
require considerable ingenuity. They present a serious technical challenge to
marine biologists because plankton have complex physiology and behaviour (Geider et al 1997). But the experimental challenges are no greater than those confronting particle physicists in search of the Higgs boson.
The task of establishing a portfolio ofbiological primitive equations to be used in
modelling plankton is also daunting because of the remarkable diversity of species
(Ormond, Gage and Angel 1997), and the complexity of their physiology and
behaviour (for an example see Geider et al. 1997). A substantial research programme will be needed to establish BPEs for the thousand most common plankton
species. Marine biologists have already taken useful steps towards that goal. For
recent reviews ofthe biology ofphytoplankton see Falkowski & Raven (1997), for
zooplankton see Harris et al. (2000) and Mauchline (1998). But, given the scale of
the task, the present rate of progress is too slow, and needs to be accelerated if BPE
modelling ofthe plankton ecosystem is to become a reality in the next few decades.
18.5 Modelling with biological primitive equations
Biological primitive equations describe the basic physiological and behavioural
functions of individual plankters. They do not describe the populations of those
organisms. Changes in populations are caused by complex interactions between
many biologic al primitive equations. It is the task of BPE modelling to compute
ples, plankton species that falI into the mindless category are responsible for toxic
algal blooms, eutrophication, cholera epidemics, and the biologic al pump in the
global carbon cycle (and therefore global warming in the 21 st century and at the
end ofice ages). Primitive equation modelling is the most promising technique for
predicting these phenomena.
18.4.4 Reproducible laboratory experiments
It will first be necessary to devise reproducible laboratory experiments to establish biological primitive equations for each of the basic functions of the physiology
and behaviour of plankton. Because these organisms are tiny, the experiments will
most likely be performed on populations rather than individuals. However, that is
not a problem ifthe individuals in the populations are identical, being clones from
a single egg, and therefore identical geneticalIy. Dr.Carlotti has drawn my attention
to the existence of significant intra-population variation in experiments with plankton from the same clutch of eggs, rather than from the same egg.
The BPEs extracted from such experiments describe aspects of the physiology
and behaviour of a single organism acting in response to its ambient environment
(which will be carefulIy controlled in the experiment). Note that the ambient environment includes physical, chemical and biological variables. The ambient biological environment experienced by a single plankter comprises the local
concentration and biologic al state of its prey, its predators, its own species and
competitors. Experiments designed to produce biological primitive equations
require considerable ingenuity. They present a serious technical challenge to
marine biologists because plankton have complex physiology and behaviour (Geider et al 1997). But the experimental challenges are no greater than those confronting particle physicists in search of the Higgs boson.
The task of establishing a portfolio ofbiological primitive equations to be used in
modelling plankton is also daunting because of the remarkable diversity of species
(Ormond, Gage and Angel 1997), and the complexity of their physiology and
behaviour (for an example see Geider et al. 1997). A substantial research programme will be needed to establish BPEs for the thousand most common plankton
species. Marine biologists have already taken useful steps towards that goal. For
recent reviews ofthe biology ofphytoplankton see Falkowski & Raven (1997), for
zooplankton see Harris et al. (2000) and Mauchline (1998). But, given the scale of
the task, the present rate of progress is too slow, and needs to be accelerated if BPE
modelling ofthe plankton ecosystem is to become a reality in the next few decades.
18.5 Modelling with biological primitive equations
Biological primitive equations describe the basic physiological and behavioural
functions of individual plankters. They do not describe the populations of those
organisms. Changes in populations are caused by complex interactions between
many biologic al primitive equations. It is the task of BPE modelling to compute