ocean: recent progress and missing gaps in the application of biogenic opal as a paleoproductivity proxy. Global and Planetary
Change, 26, 317–365.
Richaud, M., Loubere, P., Pichat, S., and Francois, R., 2007. Changes
in opal flux and the rain ratio during the last 50,000 years in the
equatorial Pacific. Deep-Sea Research Part II, 54, 762–771.
Sachs, J., and Anderson, R., 2005. Increased productivity in the subantarctic ocean during Heinrich events. Nature, 434, 1118–1121.
Sarmiento, J., and Gruber, N., 2006. Ocean Biogeochemical
Dynamics. Princeton: Princeton University Press, p. 526.
Schmiedl, G., and Mackensen, A., 1997. Late quaternary paleoproductivity and deep water circulation in the eastern South Atlantic
Ocean: evidence from benthic Foraminifera. Palaeogeography
Palaeoclimatology Palaeoecology, 130, 43–80.
Sigman, D. M., and Hain, M. P., 2012. The biological productivity
of the Ocean: section 1. Nature Education Knowledge, 3(10),
21. www.nature.com/scitable/knowledge/library/the-biologicalproductivity-of-the-ocean-section-70631104
Stoll, H., Ziveri, P., Shimizu, N., Conte, M., and Theroux, S., 2007.
Relationship between coccolith Sr/Ca ratios and coccolithophore
production and export in the Arabian Sea and Sargasso Sea.
Deep-Sea Research Part II, 54, 581–600.
Toggweiler, J., 1999. Variation of atmospheric CO 2 by ventilation
of the ocean’s deepest water. Paleoceanography, 14, 571–588,
doi:10.1029/1999PA900033.
Wassman, P., 1998. Retention versus export food chains: processes
controlling sinking loss from marine pelagic systems.
Hydrobiologia, 363, 29–57.
Zeebe, R., and Wolf-Gladrow, D., 2001. CO 2 in Seawater: Equilibrium, Kinetics, Isotopes. Amsterdam: Elsevier. Oceanography
Series, Vol. 65, p. 346.
Cross-references
Biogenic Barium
Carbonate Dissolution
Calcite Compensation Depth (CCD)
Deep-sea Sediments
Diatoms
Dust in the Ocean
Export Production
Foraminifers (Benthic)
General ocean circulation - its signals in the sediments
Marine Microfossils
Organic Matter
Paleoceanographic Proxies
Silica
PALYNOLOGY (POLLEN, SPORES, ETC.)
Anne de Vernal
Centre GEOTOP, University of Québec in Montréal,
Montréal, QC, Canada
Palynology sensu stricto and sensu lato
The word “palynology,” which is the study of pollen,
comes from the Greek word palunein (to sprinkle) in reference to the pollen that is sprinkled as dust during the
blooming season. Palynology became a scientific discipline in the early 1990s after the pioneer works of Lennart
von Post, a Swedish geologist who produced the first
pollen diagrams based on the identification and count of
pollen grains from peat deposits, to reconstruct the
postglacial vegetation of Western Europe (cf. Manten,
1967). Since then, palynology has been widely used to
reconstruct the history of vegetation through time and past
climate/environmental conditions. In botany and ecology,
palynology is often associated with the study of pollen
from flowering plants. In geosciences at large, palynology
is used in a much broader sense because palynological
slides prepared from sediment for the observation of pollen grains in optical microscopy contain many other biogenic remains. By extension, palynology thus
corresponds to the study of all microfossils composed of
highly resistant organic matter, also called sporopollenin,
which is recovered after chemical digestion of carbonate
and silica particles with hydrochloric and hydrofluoric
acid treatments, respectively.
In marine sediments, the acid-resistant organic-walled
microfossils or palynomorphs may include remains from
the planktonic organisms such as cysts of dinoflagellates
(or dinocysts), phycoma of prasinophytes, acritarchs,
tasmanids, cysts and lorica of tintinnids, copepod eggs,
etc. (e.g., Jansonius and McGregor, 1996). They may also
include organic linings of benthic foraminifers (de Vernal
et al., 1992) and allochthonous remains from freshwater
environments such as Chlorococcales (e.g., Pediastrum
spp.), in addition to pollen and spores from the terrestrial
vegetation (e.g., Traverse, 2007). Here, we focus on pollen
and spores that are particularly useful in marine
geosciences as they permit land-sea correlations in addition to the reconstruction of past vegetation and climate
in terrestrial areas from where pollen and spore assemblages originate.
Sediment sample processing for pollen analyses
The preparation of sediment samples for palynological
analyses consists in mechanical and chemical procedures
to concentrate organic-walled microfossils (e.g., Traverse,
2007; de Vernal et al., 2010). Mechanical procedures
include wet sieving for eliminating small silt size and sand
fractions and/or dense liquid separation. Chemical procedures consist in hydrochloric acid (HCl) and hydrofluoric
acid (HF) maceration to dissolve carbonate and silica particles. They may also include oxidation with potassium
hydroxide (KOH) or acetolysis, which is a treatment with
acetic anhydride ((CH 3 CO) 2 O) and sulfuric acid (H 2 SO 4 )
to eliminate labile organic matter. While oxidation techniques are often used for pollen and spore studies from
lake sediment or peat, they are avoided in marine palynology as they can alter the preservation of some marine
palynomorphs such as dinoflagellate cysts. Moreover,
the palynology of marine sediment rarely requires oxidation as the organic carbon content is generally low especially in deep-sea settings due to its degradation in the
water column and because of diagenetic processes.
Pollen and spores are usually observed, identified, and
counted with optical microscope at Â400 to Â1,000
PALYNOLOGY (POLLEN, SPORES, ETC.)
653
Change, 26, 317–365.
Richaud, M., Loubere, P., Pichat, S., and Francois, R., 2007. Changes
in opal flux and the rain ratio during the last 50,000 years in the
equatorial Pacific. Deep-Sea Research Part II, 54, 762–771.
Sachs, J., and Anderson, R., 2005. Increased productivity in the subantarctic ocean during Heinrich events. Nature, 434, 1118–1121.
Sarmiento, J., and Gruber, N., 2006. Ocean Biogeochemical
Dynamics. Princeton: Princeton University Press, p. 526.
Schmiedl, G., and Mackensen, A., 1997. Late quaternary paleoproductivity and deep water circulation in the eastern South Atlantic
Ocean: evidence from benthic Foraminifera. Palaeogeography
Palaeoclimatology Palaeoecology, 130, 43–80.
Sigman, D. M., and Hain, M. P., 2012. The biological productivity
of the Ocean: section 1. Nature Education Knowledge, 3(10),
21. www.nature.com/scitable/knowledge/library/the-biologicalproductivity-of-the-ocean-section-70631104
Stoll, H., Ziveri, P., Shimizu, N., Conte, M., and Theroux, S., 2007.
Relationship between coccolith Sr/Ca ratios and coccolithophore
production and export in the Arabian Sea and Sargasso Sea.
Deep-Sea Research Part II, 54, 581–600.
Toggweiler, J., 1999. Variation of atmospheric CO 2 by ventilation
of the ocean’s deepest water. Paleoceanography, 14, 571–588,
doi:10.1029/1999PA900033.
Wassman, P., 1998. Retention versus export food chains: processes
controlling sinking loss from marine pelagic systems.
Hydrobiologia, 363, 29–57.
Zeebe, R., and Wolf-Gladrow, D., 2001. CO 2 in Seawater: Equilibrium, Kinetics, Isotopes. Amsterdam: Elsevier. Oceanography
Series, Vol. 65, p. 346.
Cross-references
Biogenic Barium
Carbonate Dissolution
Calcite Compensation Depth (CCD)
Deep-sea Sediments
Diatoms
Dust in the Ocean
Export Production
Foraminifers (Benthic)
General ocean circulation - its signals in the sediments
Marine Microfossils
Organic Matter
Paleoceanographic Proxies
Silica
PALYNOLOGY (POLLEN, SPORES, ETC.)
Anne de Vernal
Centre GEOTOP, University of Québec in Montréal,
Montréal, QC, Canada
Palynology sensu stricto and sensu lato
The word “palynology,” which is the study of pollen,
comes from the Greek word palunein (to sprinkle) in reference to the pollen that is sprinkled as dust during the
blooming season. Palynology became a scientific discipline in the early 1990s after the pioneer works of Lennart
von Post, a Swedish geologist who produced the first
pollen diagrams based on the identification and count of
pollen grains from peat deposits, to reconstruct the
postglacial vegetation of Western Europe (cf. Manten,
1967). Since then, palynology has been widely used to
reconstruct the history of vegetation through time and past
climate/environmental conditions. In botany and ecology,
palynology is often associated with the study of pollen
from flowering plants. In geosciences at large, palynology
is used in a much broader sense because palynological
slides prepared from sediment for the observation of pollen grains in optical microscopy contain many other biogenic remains. By extension, palynology thus
corresponds to the study of all microfossils composed of
highly resistant organic matter, also called sporopollenin,
which is recovered after chemical digestion of carbonate
and silica particles with hydrochloric and hydrofluoric
acid treatments, respectively.
In marine sediments, the acid-resistant organic-walled
microfossils or palynomorphs may include remains from
the planktonic organisms such as cysts of dinoflagellates
(or dinocysts), phycoma of prasinophytes, acritarchs,
tasmanids, cysts and lorica of tintinnids, copepod eggs,
etc. (e.g., Jansonius and McGregor, 1996). They may also
include organic linings of benthic foraminifers (de Vernal
et al., 1992) and allochthonous remains from freshwater
environments such as Chlorococcales (e.g., Pediastrum
spp.), in addition to pollen and spores from the terrestrial
vegetation (e.g., Traverse, 2007). Here, we focus on pollen
and spores that are particularly useful in marine
geosciences as they permit land-sea correlations in addition to the reconstruction of past vegetation and climate
in terrestrial areas from where pollen and spore assemblages originate.
Sediment sample processing for pollen analyses
The preparation of sediment samples for palynological
analyses consists in mechanical and chemical procedures
to concentrate organic-walled microfossils (e.g., Traverse,
2007; de Vernal et al., 2010). Mechanical procedures
include wet sieving for eliminating small silt size and sand
fractions and/or dense liquid separation. Chemical procedures consist in hydrochloric acid (HCl) and hydrofluoric
acid (HF) maceration to dissolve carbonate and silica particles. They may also include oxidation with potassium
hydroxide (KOH) or acetolysis, which is a treatment with
acetic anhydride ((CH 3 CO) 2 O) and sulfuric acid (H 2 SO 4 )
to eliminate labile organic matter. While oxidation techniques are often used for pollen and spore studies from
lake sediment or peat, they are avoided in marine palynology as they can alter the preservation of some marine
palynomorphs such as dinoflagellate cysts. Moreover,
the palynology of marine sediment rarely requires oxidation as the organic carbon content is generally low especially in deep-sea settings due to its degradation in the
water column and because of diagenetic processes.
Pollen and spores are usually observed, identified, and
counted with optical microscope at Â400 to Â1,000
PALYNOLOGY (POLLEN, SPORES, ETC.)
653