Produccion biologica y oceanografa en giros centrales

From: M Blackburn 1981. In: Analysis of marine ecosystems (A.R.Longhurst, ed.), Academic Press, pp.3-29

Photograph: Richa Sharma

Principales caractersticas
500 m profundidad (P yA) Acompaados pequeos giros Escalas 1000km (horizontal) y centenas m (vertical) Se consideran un ecosistema ocenico Formados por 4 sitios: Pacifico (N y S), Atlntico (N y S), Indico (S) y el periodo de inversin en la circulacin superficial debido a eventos Monsnicos. Se diferencian de procesos de Nucleo frio y clido de la Corriente kuroshio y Golfo.

Composicin Giros
Dos en Pacfico, dos Atlntico y zona sur del Indico. Cada giro se compone por una corriente Ecuatorial dirigida al occidente, una corriente de margen occidental dirigida hacia el polo, una corriente latitud media (40L) dirigida hacia el Este (CDO) y una corriente de margen oriental dirigida hacia el Ecuador. Visin antigua era simplista

Anlisis topografa dinmica anual del mar superficial

La carta revela una elevacin entre 20 y 25 Lat Norte en el Pacfico Occidental (Yoshida & Kidoro, 1967). Hoy se sabe que se compone de3 giros menores. Existe una corriente hacia el Este a app 10 S.
Worthington (1976) distingui dos claros anticiclones en el Atlntico Norte (40 y 50 LN). Zona occidental Pacfico sur se compone varios giros anticiclnicos. Conclusin: Giros anticilnicos son ms complejos que lo pensado originalmente

Caractersticas fsico-qumicas de los giros importantes

- Son clidos Aguas de alta transparencia (100 m) Son convergentes Velocidades superficiales corrientes: 5 a 10 cm/s Capa mezcla superior 15 a120 m prof. Termoclina 100 a 200 m de grosor Tienen alta salinidad (> 35PSU) Alto contenido OD Baja concentracin de P y N (Paradoja del Plancton) La distribucin horizontal es homognea excepto en los mrgenes T, S, y OD son mas altos en el margen occidental. Nutrientes ms bajos en el margen occidental. Cambios estacionales leves a moderados. Se consideran las zonas de los ocanos biolgicamente mas oligotrficas.

Burkov & Monin (1973)

Variable
Capa Mezcla (m, V) Capa Mezcla (m, I)

Giro Subpolar
15-30 > 120

Divergencia Ecuatorial
25-65 25-65

Giro subtropical
15-50 65-120

Capa Euftica (m)

NO3 (100 m) ug-at l-1 Produccin Primaria (g C m-2 da-1)

20-50
10-25 0,1-0,5

45-85
10-25 0,1-0,5

75-150
0-5 < 0,1

Clorofila a (mg m-2)

Biomasa X herbvoros (ug C) Zooplancton (g m-2)*

15-150
40-400 30-50

15-30
4-40 7-25

5-25
4-40 <7

Zooplancton (g m-2)**
Micronecton (g m-2)* Micronecton (g m-2)** Bentos (g m-2)

150
0,6 2 0,1-1,0

8-13
0,6-0,8 1-3 0,1-1,0

9
0,3 1 <0,05

PP y Chl a integrada para capa euftica

(*) 0- 200 m; (**) 0-1000 m

Fuente: Longhurst (1981)

[Fe] =

Caractersticas Plancton: tamao

Tamao (um) < 0,2 0,2-2,0 2,0-20,0 20,0-200 200-20000 (20 mm) Ejemplos Femtoplankton (virus) Picoplankton (bacterias y eucariotas) pequeos Nanoplankton (diatomeas, dinoflagelados, radiolarios) Microplankton (diatomeas, dinoflagelados, protozoos, nauplios copepodos) Mesoplankton (mayoria zooplankton)

2-20 cm

Macroplankton

Plancton: organismos que derivan (gr.: planktos)

Tramas troficas en ambientes oceanicos oligotroficos

Modelo Sverdrup (1953) Porque existen blooms y Como estimar PP

Tasas crecimiento fito = 1/t ln (N/No) = 1/t log2 (N/No) Unidades: div/da

PP e intensidad luminica

CURVAS P/I = PENDIENTE P MAX = Tasa maxima fijacion carbono I = intensidad luminica al inicio de la fotoinhibicion (Trevor Platt et al., 1980)

Modelo Prof critica Sverdrup (1953)

Recordatorio Fotosintesis

http://www.jochemnet.de/fiu/OCB3043_22.html

Modelo Prof. Crtica Sverdrup (1953)

1. Photosynthesis decreases exponentially with depth due to decrease in light availability 2. Respiration is unaffected by light and remains constant with depth. 3. Phytoplankton is mixed by turbulence and experiences different light intensities over time, sometimes above and sometimes below compensation point 4. Critical depth = depth at which photosynthesis of the total water column phytoplankton population equals their total respiration

De que depende la PP
Especie fitoplanctnica Edad especie fitoplanctnica Grupo taxonmico (Dinoflagelados v/s diatomeas) Intensidad luminica Transparencia agua Hora del da Estacin climtica Disponibilidad nutrientes (P, N, Si, Fe) Temperatura Abundancia pastoreadores o herbvoros Profundidad mezcla Zona ocano Concentracin inicial o tamao parche Potencial dispersin

Tipos Fitoplancton

Variabilidad mesoescala-Tamao fitoplankton

- 1 = fitoplancton pequeo + 1= fitoplancton grande

(a) biomasa, (b) produccin primaria (c) tasa crecimiento especifico. Escala espacial: 256 x 256 km

Martin et al 2001 J Mar System

Variacion espacial y temporal Fitoplancton (das)

Importancia PP

http://earthobservatory.nasa.gov

Fitoplancton y cambios ambientales

Modelos qumicos y biologicos predicen que el calentamiento de la superficie del mar (greenhouse gases) implicara una declinacin fitoplancton. Esto implicara una caida en la produccion primaria dado que la columna de gua se estratificara y no permitira el ascenso nutrientes o el reciclaje de los nutrientes.

PP a escala global

http://earthobservatory.nasa.gov

Ciclo N2

Ciclo N coastal shelf and upwelling (I), OMZs (II), surface waters of the open ocean (III), and deep water (IV). PON, particulate organic nitrogen. Pathways: A, DIN assimilation; B, ammonium regeneration; C, nitrogen fixation; D, nitrate diffusion/advection from deep water; E, nitrification; F, nitrification; G, denitrification.

1953
Google citation count: 561

Net phytoplankton growth occurs when area acd > abdf

 Sverdrups 1953 paper was a formalization of the critical depth concept originally proposed by Gran and Braarud in 1935*

The critical depth hypothesis attempts to explain what initiates a vernal bloom, not what controls the magnitude of a bloom
A bloom is an increase in biomass, not photosynthetic rate

The hypothesis states that a bloom begins when the mixed layer shoals to a depth above the critical depth horizon where production (P) > respiration (R)
R = grazing + sinking + phytoplankton respiration + all other losses R is assumed constant Inverse of Sverdrup: prior to crossing the critical depth criterion, net growth is negligible or negative

* Gran & Braarud. 1935. J. Biol. Board Can. 1 (5), 279-467

 Sverdrup: net growth can be independent of gross production under heavy grazing Sverdrup: the bloom observed 2 days after the depth of the mixed layer was for the first time smaller than the critical depth likely reflected advection not rapid local growth Sverdrup: the first increase in biomass occurred before stratification Sverdrup: It is therefore not advisable to place too great emphasis on the agreement between theory and [the Weather Ship M] observations (occurrence of blooms in the absence of stratification is not uncommon)

* e.g., Townsend et al. 1992. Nature 360, 59-62

Abandoning Sverdrup
80 70
NA-12

70

60

50

40

30

20

10

A
NA-11 NA-9 NA-8 NA-5 NA-2 NA-10 NA-7 NA-4 NA-1

70

60

60

50

NA-6 NA-3

50

40

40

30
0.01 0.1 1.0 10

30

Chlorophyll (mg m-3)

SeaWiFS data 1998 2006 8-day resolution 12 central NA bins, minimize advect. Chlsat = OC4-V4 Cphyto = GSM / Westberry et al 2008

= 1989 NABE = 2008 NAB

= focus in later slides

Cphyto (mg C m-3 )

80 60 40 20 0

NA-5 - Latitude range: 45o 50oN

1.6 1.4 1.2 1.0 0.8 0.6 0.4 0.2 0.0

Chlsat (mg m-3 )

1998

1999

2000

2001

2002

2003

2004

2005

2006

Year
North Atlantic seasonal cycles are dominated by changes in biomass Thus, Cphyto ~ Chlsat Differences between Cphyto and Chlsat consistent with photoacclimation All analyses have been completed with both C and Chl Results to follow are the same irrespective of C or Chl Remaining slides focus on C within-bin standard deviations shown above

Cphyto (mg C m-3 )

80 60 40 20 0

NA-5 - Latitude range: 45o 50oN

1998

1999

2000

2001

2002

2003

2004

2005

2006

PAR

Year

MLD

peak biomass occurs in spring coincident with rising PAR and shoaling MLD also associated with rapid rise in primary production Conclusion: phytoplankton in the North Atlantic exhibit a repeated vernal bloom caused by increased primary production and growth associated with rising light and shallowing mixed layers aka, Sverdrup
unfortunately, biomass can be a terribly misleading thing. & correlation is not causation

Bloom in a Bottle
-3 Chlorophyll concentration m ) Chlorophyll (mg concentration (mg m )

To understand what causes a bloom, it is necessary to first identify when a bloom started
12 10 8 6 4 2 0
0 2 4 6 8 10

The start of a bloom can not be defined by biomass - e.g., when biomass X mg m-3 or Y% above annual median
5% > mean

-3

Using biomass can lead to the wrong start date and association of bloom initiation with the wrong environmental forcing Bloom initiation implies a change in the rate of growth for Sverdrup it was the beginning of positive net growth An easy way to get a first-order sense of rate changes is to plot biomass on a logarithmic scale

10

0.1 0 2 4 6 8 10

perturbation

?
Days

Net growth rate = r = ln(C1/C0) = slope of log plot t1 t0

0 30 20

NA-2

-50 -100 -150 -200

Abandoning Sverdrup
The North Atlantic bloom does not begin in the spring Net exponential growth begins mid-winter Shift from negative to positive biomass changes coincides with the cessation of mixed layer deepening Net growth rates are, on average, comparable from winter through spring

10 9 8 7 40 30 20

-250 -300 0

NA-5

-50 -100 -150

Cphyto (mg C m )

10 9 8 7 6 50 40 30 20 10 9 8 7 6 5 4 80 60 40 20

-200 -250 -300 0

-3

MLD (m)

NA-8

-50 -100 -150 -200 -250 -300 0

Net growth rates do not reflect changes in incident light, photosynthetic rate, or gross growth rate ()
The critical depth hypothesis can be dismissed
100 80 60 40
NA-5 - Latitude range: 45o 50oN

NA-11
-100 -200 -300

Cphyto

20 10 8 6 4 2

10 8 6
J F MAM J J A S ON D

-400 -500

Month

1998

1999

2000

2001

2002

2003

2004

2005

2006

Year

Bloom in a Bottle
60 1.0 0.8 0.6 20 0.4 0 -50

Cphyto (mg m )

-3

40

-100

-150 -200 -250 -300

Population specific net growth rates (r) can be calculated from changes in phytoplankton concentration (m-3) so long as the mixed layer is either shoaling or not deepening
r = ln(C1/C0) t1 t0

Chlsat (mg m )

MLD (m)

10 8 6 J F M A M J J A S O N D

-3

0.2

However, one must consider the influence of dilution when the mixed layer is deepening
r = ln[(C1 MLD1) /[(C0MLD0)] t1 t 0

A dilution correction should be considered when assessing growth rates during mixed layer deepening

NA-5 - Latitude range: 45o 50oN

Deep

Shallow

Population net specific growth rate (r) for the active water column becomes positive in late-autumn / early winter and remains positive through the spring until nutrients are depleted

Growth-phase maxima in r can occur during MLD deepening, MLD maximum, or MLD shoaling
Overall, r is inversely related to PAR and In 100% of available complete annual cycles, r becomes positive before PAR begins to increase

NA-5 - Latitude range: 45o 50oN

Shallow

Deep Starting now in July

The vernal bloom appears to be an event initiated in late fall Triggering of the bloom appears to be associated with mixed layer deepening (not shoaling) How is this possible? Why the mid-winter decrease in r?

How is this possible?

NA-5 - Latitude range: 45o 50oN

* = NPP / CZ

*
: (d-1) r (d )
-1

1.0

0.8

0.6

0.4

0.2

0.0 J A S O N D J F M A M J J

Month

A net specific growth rate of 0.02 implies approximately 1 division per month Typical winter C = 4 8 mg m-3, Typical spring C peak = 25 70 mg m-3 NA bloom requires 2 4 doublings over 3 - 4 months, or average r of 0.009 to 0.03 d-1

1.0

10000

: (d-1)

-2

0.8

"Respiration" (mg C m d )

-1
0.6 0.4

1000

r (d )

100

-1

0.2

0.0 J A S O N D J F M A M J J

10 10 100 1000 10000

Month

Production (mg C m-2 d-1)

Positive r through winter is allowed because losses co-vary with (Sverdrup assumed this respiration to be a constant) r 0 as PAR 0 at very high latitudes in mid-winter (no light) a critical depth can never be reached The increase in r during winter implies that the fraction of that escapes predation and other losses (i.e., r:) must increase in the winter

The mid-winter decrease

Diluted Digression

Landry & Hassett 1982 Mar. Biol. 67, 283-288 Landry et al. 1995 Mar. Ecol. Prog. Ser. 120, 53-63

The Grand Dilution Hypothesis

* As a replacement for the Critical Depth Hypothesis, it is proposed that the north Atlantic bloom is a consequence of a massive scale dilution experiment Mixed layer deepening causes a slight decoupling between phytoplankton growth and losses (grazing, mostly)

0.20

0 -50

The decoupling increases so long as the mixed layer continues to deepen

Mixed layer shoaling drives a re-coupling of phytoplankton growth and losses (grazing) Thus, while spring shoaling and increasing light favor enhanced photosynthesis and growth, they also favor heavier grazing losses

0.15

-100

MLD & Zeu (m)

r::

0.10

-150 -200 -250 -300 J A S O N D J F M A M J J

0.05

0.00

Month Landry & Hassett 1982 Mar. Biol. 67, 283-288 Landry et al. 1995 Mar. Ecol. Prog. Ser. 120, 53-63

Modeling the Grand Dilution

As an initial attempt, a simple model was developed and compared to average annual cycles of the r: ratio and r for each of the 12 bins Model input was the value of r: during the first week in July (-0.01) and MLD and Zeu (from Chlsat) The 3 model conditions were as follows:

MLD0

Mixed layer deepening within the euphotic zone, entrains phytoplankton and grazers

NO CHANGE in r:
Zeu

Modeling the Grand Dilution

As an initial attempt, a simple model was developed and compared to average annual cycles of the r: ratio and r for each of the 12 bins Model input was the value of r: during the first week in July (-0.01) and MLD and Zeu (from Chlsat) The 3 model conditions were as follows:

Zeu

Mixed layer deepening below euphotic zone, entrains phytoplankton free water Dilutes predators & Prey

MLD0

r: CHANGES IN PROPORTION TO DILUTION

Modeling the Grand Dilution

As an initial attempt, a simple model was developed and compared to average annual cycles of the r: ratio and r for each of the 12 bins Model input was the value of r: during the first week in July (-0.01) and MLD and Zeu (from Chlsat) The 3 model conditions were as follows:

Zeu

Mixed layer shoaling cuts off the lower population of phytoplankton, has no direct effect on phytoplankton concentration, but concentrates mobile grazers Shoaling concentrates predators but not prey

r: CHANGES IN PROPORTION TO MLD CHANGE BUT AT A SLOWER RATE THAN DEEPENING EFFECT
MLD0

Modeling the Grand Dilution

Final Comments
Temporal coverage of the satellite record provides a unique opportunity to reevaluate bloom dynamics The critical depth hypothesis is found wanting (actually, it fails miserably) A Grand Dilution Hypothesis is suggested, but is not the only potential explanation (aggregation, temperature effects, sinking.?) Dilution Hypothesis accommodates blooms w/o stratification

Climate change effects on North Atlantic (and other) blooms may be very much different for a Critical Depth concept of blooms and a Dilution concept of blooms
Revisiting bloom experimentation in North Atlantic?


Winter Chlz Lat trends

Mixing velocity
Feb C max Lat r

Cphyt vs Csat

GIROS CENTRALES DE LOS OCEANOS

NICHO (JOSEPH GRINELL, 1917) CONDICIONES DEL HABITAT QUE PERMITEN A UNA ESPECIE EXISTIR

NICHO MULTIDIMENSIONAL (G. EVELYN HUTCHINSON, 1957) MUCHAS VARIABLES DEL AMBIENTE RESTRINGEN LA EXISTENCIA DE UNA ESPECIE

NICHO REALIZADO V/S NICHO FUNDAMENTAL (HUTCHINSON, 1957)

CHARLES ELTON (1927) UNA ESPECIE PODRIA OCUPAR MAS DE UN NICHO UN NICHO PODRIA SER OCUPADO POR MULTIPLES ESPECIES LAS COMUNIDADES PUEDEN TENER NICHOS VACIOS NO OCUPADOS POR ESPECIES.

METAPOBLACIONES Y DISPERSION REFINAMIENTO HIPOTESIS ANTERIORES POBLACIONES FORMAN REDES INTERACTIVAS O METAPOBLACIONES, PERMITIENDO EL FLUJO GENICO Y DE INDIVIDUOS ENTRE DISTINTAS POBLACIONES UN INDIVIDUO PUEDE SOBREVIVIR FUERA DEL NICHO Y NO MANTENER LA VIABILIDAD DE LA POBLACION APARECE EL CONCEPTO DE POBLACIONES FUENTES Y POBLACIONES RECEPTORAS

EN ESTE CASO, EL NICHO REALIZADO PUEDE SER MAYOR QUE EL NICHO FUNDAMENTAL

HIJO/AS PUTATIVOS DE EVELYN HUTCHINSON

G. Evelyn Hutchinson,1961
Porque existen numerosas especies filogenetica y funcionalmente similares en zonas de baja PP Como logran coexistir? Es fundamental para comprender como se conserva la biodiversidad en zonas oligotroficas de los ocanos Pueden coexistir especies que dependen de un mismo recurso Puede haber exclusion competitiva o segregacin nichos? Esta se reconoce como la paradoja del plancton propuesta por una de los padres dela Ecologa Moderna

Mas Plankton paradox

Cuando se intenta explicar el origen y la conservacin de la diversidad de la vida en la Tierra, se observa que existen dos escuelas de pensamiento: 1) Teora Nicho: Especies coexisten porque son distintas.

2) Teora Neutral: Especies coexisten porque son idnticas.

La existencia de diferencias entre especies ha sido considerada como evidencia irrefutable que capacita a las especies a coexistir. Sin embargo, recientes trabajos han identificado una clase de modelos que se basan en que las especies coexisten porque son idnticas.

Sin embargo, Modelos Neutrales generan y explican importantes patrones biolgicos de variacin tales como la distribucin espacial de especies y riqueza de especies utilizando tres parmetros: el tamao de la metacomunidad, estrategias de dispersin y tasas de especiacin.
Verdades Cientficas: 1) La verdad probablemente se encuentre entre ambos tipos de teoras. 2) El modelo no es nuevo. Al contrario, es uno de los ms antiguos conceptos ecolgicos: modelo competencia de Lovtka y Volterra. (Teora neutralidad emergente: complementa ambos tipos de teoras. Si bien es un modelo fundamentalmente basado en el concepto de nicho, el uso de coeficientes de competitividad generan patrones neutrales a partir de las simulaciones que involucran gran nmero de competidores).

La paradoja del Plancton: Evidencias ecolgicas y diversidad

If all phytoplankton species require essentially the same resources, how can so many species co-exist within the same lake? ("paradox of the plankton") Hutchinson (1961) Homage to Santa Rosalia

Paradoja Plancton (Hutchinson 1961): Molecular genetic evidences in marine microalgae

Ostreococcus
Eukaryotic phytoplankton exhibit great diversity that contrasts with the lower apparent diversity of ecological niches available to them in aquatic ecosystems as Gyral. Know as the paradox of the plankton, has long puzzled biologists (2). By providing molecular level information on related species, genomics is poised to provide new insights into this paradox. Picophytoplankton, with cell diameters <2 m, play a significant role in major biogeochemical processes, primary productivity, and food webs, especially in oligotrophic waters. Within this size class, the smallest known eukaryotes are Ostreococcus tauri and related species. Although more similar to flattened spheres in shape, these organisms are 1 m in diameter (3, 4) and have been isolated or detected from samples of diverse geographical origins (58). They belong to the Prasinophyceae, an early diverging class within the green plant lineage, and have a strikingly simple cellular organization, with no cell wall or flagella, and with a single chloroplast and mitochondrion (4). Recent work has shown that small-subunit rDNA sequences of Ostreococcus from cultures and environmental samples cluster into four different clades that are likely distinct enough to represent different species (6, 9). Here we report on the gene content, genome organization, and deduced metabolic capacity of the complete genome of Ostreococcus sp. strain CCE9901 (7), a representative of surfaceocean adapted Ostreococcus, referred to here as Ostreococcus lucimarinus. We compare it to the analogous features of the related species O. tauri strain OTH95 (10). Our results show that many processes have been involved in the evolution and speciation of even these sister organisms, from dramatic changes in genome structure to significant differences in metabolic capabilities.

Ostreococcus Clase Prasinophyceae

Es un gnero de microalga unicelular verde de aspecto cocoidal. Fue descubierta en 1994 rasinophyceae. It was discovered in 1994 in the Lagoon Thau,desembocadura ro Rhone por Courtiesy se ha descubierto en muchas zonas ocenicas. Es la ms pequea celula eucariotaconocida con untamao de1 um. Su genoma nuclear tiene 12.56 Mb. Ostreococcus tauri tiene un nucleo con 14 cromosomas lineales,un cloroplasto y algunas mitocondrias.

Multidimensionalidad
Clarke et al 2007

Textural Holling
Holling 1992

Neutralidad emergente Schaeffer & van Ness 2006

Neutralidad

Hubbells (Hubbell 2001) T Neutral

T Nicho

Ecological Letters 2009

Explicacin de los 4 modelos

1. Multidimensionalidad (Clarke et al 2007):

2.

Modelo Textural Holling (Holling, 1992)

3.

Modelo Neutralidad emergente (Scheffer & van Ness, 2006): predice agregados de especies alo largo de un eje nico del nicho. Incluye aspectos de la teoria del nicho y neutral.

Explicacin de los 4 modelos

Hiptesis neutralidad Hubbells (2001)

Se proponen 3 predicciones:

Se plantean 4 predicciones

Prediccion 3

Prediccion 4

Professor Celso Grebogi

Are fractal skeletons the explanation for the plankton paradox and narrowing of arteries due to cell trapping in a disturbed blood flow? Institute for Complex Systems, King's College, University of Aberdeen, Scotland, UK Professor Grebogi researched in the field of plasma physics prior to his work on chaotic dynamical systems which combines analytical methods and techniques with extensive computer experiments utilising computational facilities. Important results include the establishment of crises as the fundamental process by which chaotic attractors undergo sudden changes as a system parameter varies; the mathematical theory and experimental verification of how transient chaos phenomena are likely to manifest themselves in practice; and the work on fractal basin boundaries. Current research focuses on methods to control chaos, the dynamics of spatio-temporal systems, active processes in chaotic flows, and the rigorous determination of how long actual trajectories of a chaotic process stay near a given numerical trajectory, i.e. the problem of shadowing.

Paradoja del Plancton a nivel larval

Larvas con alto potencial dispersion geografica

Paradoja del Plancton a nivel Holoplancton

Segregacion nicho: escala vertical copepodos

Anatomia copepodo

Alimentacin copepodos
Poseen dos velocidades de natacin. El primero es lento, constante y realizado con sus piezas bucales. La segunda es una sucesin de saltos separados por la quietud. Este tipo de natacin lo logran los apndices traxicos. Las antenas concentran y seleccionan partculas.

Estadios quiescentes o dormantes Neocalanus tonsus (aguas oceanicas)

Nauplios copepodos y cirripedios

Copepoda

Cirripedia

Copepodos y cambio climatico

Escalas variabilidad espacial en el Oceano Pacifico

Variabilidad temporal

Corriente Kuroshio y dinamica zooplancton

Ocean Production

1. World patterns of primary production 2. Dominant communities of the North Pacific 3. North American coastal communities

Percent of World's Surface Area by Bioclimatic Region

Polar 2% Boreal 5% Subboreal 5% Subtropical 5% Ocean 72% Tropical 11% Freshwater 0.1%

Percent of World's Annual Primary Production

Freshwater 0.01% Ocean 26% Tropical 43% Polar 1% Boreal 7% Subboreal 8% Subtropical 15%

Gross Primary Production in Marine and Terrestrial Ecosystems

Ocean Area Production Land Area (g C/m2/year) Open 50-160 Deserts, Ocean grassland Production (g C/m2/year) 50

Coastal Ocean Upwelling zones

Salt marshes

100-250

Forest, crops, 25-150 pastures Rain forest, agriculture 150-500

200-800
1000-4000

Smith, S. and Hollibaugh, J., Rev. Geophys., 31, 75, 1993.

Relative Ocean Primary Production by Province

300

g C/m2/yr

200 100 0 Open Ocean Coastal Province Upwelling

Global Primary Production in Marine Ecosystems

Area Primary Production
(g C/m2/year)

World Ocean Area km2 %

Total Primary Production

(metric tons carbon/year)

Upwelling Coasts Open Ocean

640 160 50

0.36 x 106 0.1 54 x 106

0.23 x 109

15.0 8.6 x 109

307 x 106 85.0 15.3 x 109

Smith, S. and Hollibaugh, J., Rev. Geophys., 31, 75, 1993.

Annual marine primary production in the worlds oceans in relation to the area where production occurs (Longhurst et al. 1995)
Most of the ocean area is very unproductive; only a small fraction is highly productive

The proportion of ocean area by depth. The mean depth of the oceans is 3.7 km and they cover 72% of the globe.
(Sunderman 1986) 0 1

Depth (km)

Thus the great majority of the ocean is very deep

2 3 4 5 6 7 8 0 5 10 15 20 25 30 Percentage of ocean area 35

Ocean Temperatures

Relationship between depth, phytoplankton photosynthesis and phytoplankton respiration

Phytoplankton need light, nutrients and carbon dioxide to grow (nitrates, phosphates and iron can limit primary production). Phytoplankton photosynthesis Photosynthesis depends strongly on light, which declines with depth. Above the compensation depth, more carbon is fixed by photosynthesis than is lost in respiration. Below that depth (which can be as deep as 100 m or as shallow as 1-2 m) more carbon is lost than is fixed. Below the critical depth there is insufficient light for phytoplankton.

Depth

compensation depth

Phytoplankton respiration critical depth

From Sverdrup (1953)

Primary production: Enrichment Retention Concentration

Chlorophyll production, September 1998 Wide band of production in north Southern blooms just beginning Notice upwelling along west coasts of continents

(Picture courtesy of NASA SEAWIFS program)

Seasonal cycles of primary and secondary production vary among regions

phytoplankton zooplankton Arctic Ocean Biomass

Tropical Pacific Biomass J F M A M J J A S O N D Month

Temperate North Atlantic

Biomass

Cushing 1975

Relationship between annual phytoplankton production and the production of carnivorous fishes in open ocean and coastal environments
Iverson (1990, in Jennings et al.)
Fish and squid production (g/ m sq/ year) 25

15

20

NW Atlantic shelf

10

Baltic and nearby seas

Open ocean sites

0 0 50 100 150 200 250 300 Phytoplankton production (g C/m sq/ yr)

General circulation patterns

North Pacific Ocean current and gyral systems (Dodimead et al. 1963)

Catches along 155oW, July 15-30 1984

Subarctic Ridge Domain salmonids, salmon shark, spiny dogfish Transition Domain nail squid, saury, blue shark, albacore, neon flying squid Sub-tropic tuna, striped marlin, mackerel, dolphin, yellowtail,

Catches of fishes and squids in gill nets along 155oW by Oshoro Maru, 1984
Salmonids
70% 60%

Pacific Pomfret Blue Shark, Albacore,

Ommastrephes bartrami
Tuna, Marlin, Shortbill Spearfish, Mackerel, Dolphin, Yellowtail

Percent of Total

50% 40% 30% 20% 10% 0%

n = 2897 n = 694

n = 199

n = 3999

55 54 53 52 51 50 49 48 47 45 44 44 42 41 39 38 36 Latitude

Species distribution in relationship to sea surface temperature (Pearcy et al.)

Dominant epipelagic species in the Tropical Province

Wahoo (Acanthocybium solandri)

Flying fish

Yellowfin tuna (Thunnus albacares)

Tropical epipelagic species

Swordfish (Xiphias gladius)

Striped marlin (Tetrapturus audax)

Indo-Pacific sailfish (Istiophorus platypterus)

Dominant epipelagic species in Subtropical Province

Bigeye tuna (Thunnus obesus)

Dolphin (Coryphaena hippurus)

Bluefin tuna (Thunnus thynnus)

Dominant epipelagic species in Transition Province of North Pacific

Blue shark (Prionace glauca) Pacific mackerel (Scomber japanicus) Albacore (Thunnus alaunga)

Pacific pomfret (Brama japonicus)

Jack mackerel (Trachurus symmetricus)

Saury (Cololabis saira)

Generalized foodweb in the Transition Province of the North Pacific Ocean

Myctophids: lanternfish, vertically migrating meso-pelagic species

Pteropods and heteropods are planktonic molluscs

Approximate areas of oceanic domains and prevailing current directions in the Northeast Pacific Ocean (Ware and McFarlane 1989)

Ekman flow
Water velocity decreases and rotates in direction with increasing depth: The water does not go in the same direction as the wind.

Summer winds tend to come from the north, causing Ekman transport of water offshore along the surface. This brings colder, nutrient-rich water from along the continental slope to replace it. The strength of this upwelling varies from year to year.
http://pfeg.noaa.gov

Dominant species of the Coastal Upwelling Domain off the North American west coast
Pacific hake (Merluccius productus)

Pacific sardine (Sardinops sagax)

Pacific mackerel (Scomber japonicus) Northern anchovy (Engraulis mordax)

Concentrations of dominant species in the coastal upwelling domain; summer

Pacific hake

Pacific sardine

Dominant species of the Coastal Downwelling Domain off North American west coast
Pacific cod (Gadus macrocephalus) Sablefish (Anoplopoma fimbria)

Pacific halibut (Hippoglossus stenolepis)

Walleye pollock (Theragra chalcogramma)

Pacific herring (Clupea pallasi)

Dominant epipelagic fishes in the Subarctic Province of the North Pacific Ocean
Salmon shark, Lamna ditropis

Chinook salmon (O. tshawytscha)

Sockeye salmon (O. nerka)

Primary Production for North Pacific Domains (Ware and McFarlane 1989)
Domain Coastal Upwelling Primary Production (g C/m2/yr) 195 g 152 g Region California WA - OR

Coastal Downwelling
Central Subarctic

72-100 g
185-330 g 45-72 100

Oceanic
Shelf (Different estimation methods)

These regions also differ in the size and species composition, seasonal abundance patterns, and depth distributions of phyto- and zooplankton.

Biomass and potential yield for North Pacific domains

Domain
Coastal Upwelling Coastal Downwelling Central Subarctic*

Biomass (T/km2) 13.7

10.4

Yield (T/km2/yr) 3.3

1.4

Catch: Biomass 0.24

0.13

0.15

0.05

0.33

*Does not include pomfret, saury, jack mackerel and albacore tuna
Ware and McFarlane (1989)

Estimates of production and biomass of demersal fishes at different depths in the eastern North Atlantic
biomass
3 Production (g/m2/year)
Abyssal Plain

Biomass (g/m2)

production 1 0
200-600

600-1000

1-2000

2-3000

3-4000

Depth zone

Concentrations of dominant species in the coastal upwelling domain; summer

Pacific makerel

Northern anchovy

Concentration of the dominate species in the Coastal Downwelling domain

Concentration of walleye pollock in the Coastal Downwelling domain

Biomass of Salmonids
Biomass (T x 1000)

600 400 200 0 Coastal shelf Coastal downwelling Coastal upwelling

Domain

Biomass of all species

Biomass (T x 1000)
5000 4000 3000 2000 1000 0 Coastal shelf Coastal downwelling Coastal upwelling

Domain

Temperatures at 0, 50, 200 and 600 m depth

Sea Surface Temperature

Circulacin Giro Indico

Efecto monsones y GOI en sistemas locales

Ocean productivity changes dramatically in this area with the reversal of the monsoon. The CZCS Fig. 13.3 shows high productivity (red, orange and yellow) along the southwest coast of Oman, in the Persian Gulf, along the west coast of India and the east coast of Somalia.

(Jul-Sept 1979)

(Abr-Jul 1979)

Rojo, Naranja,Amarillo

= alta productividad

Azul y violeta

= Baja Productividad

Masas agua ocano Indico Diagrama T/S

Fig. 13.10 Mean TS cuves for the Indian Ocean

Circulacin mar Rojo y cua de sal

Fig. 13.11 Schematic Red Sea circulation

ENSO in the Indian Ocean

13.1.1 Indonesian Throughflow

En 1994, Tourre and White encontraron que el OI como el OP muestran seales de efectos del Nio. Como efecto de este evento una masa de agua clida se mueve hacia el Este al OI en un ciclo de 3 a7 aos. Esta masa de agua clida se mantiene 12 a 18 meses despus de ocurrido el Nio.

Flow from the Pacific to the Indian Ocean, via the complicated series of passages and sills that make up the Indonesian Archipelago (Fig. 13.8), is a oneway movement of significant amounts of freshwater and heat. The "Indonesian throughflow" has been linked to the regional climate of the Australasian region and to the large-scale structure of the global thermohaline circulation.

Fig. 13.8 Indonesian Archipelago; lines indicate the 200, 1,000 and 2,000 m isobaths and the red dots indicate the location of shallow pressure gauges