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Hydrostatic pressure substantially reduces deep-sea microbial activity

Posted by mmaheigan

· Thursday, May 11th, 2023

Deep sea microbial communities are experiencing increasing hydrostatic pressure with depth. It is known that some deep sea microbes require high hydrostatic pressure for growth, but most measurements of deep-sea microbial activity have been performed under atmospheric pressure conditions.

In a recent paper published in Nature Geoscience, the authors used a new device coined ‘In Situ Microbial Incubator’ (ISMI) to determine prokaryotic heterotrophic activity under in situ conditions. They compared microbial activity in situ with activity under atmospheric pressure at 27 stations from 175 to 4000 m depths in the Atlantic, Pacific, and the Southern Ocean. The bulk of heterotrophic activity under in situ pressure is always lower than under atmospheric pressure conditions and is increasingly inhibited with increasing hydrostatic pressure. Single-cell analysis revealed that deep sea prokaryotic communities consist of a small fraction of pressure-loving (piezophilic) microbes while the vast majority is pressure-insensitive (piezotolerant). Surprisingly, the piezosensitive fraction (~10% of the total community) responds with a more than 100-fold increase of activity upon depressurization. In the microbe proteomes, the authors uncovered taxonomically characteristic survival strategies in meso- and bathypelagic waters. These findings indicate that the overall heterotrophic microbial activity in the deep sea is substantially lower than previously assumed, which implies major impacts on the carbon budget of the ocean’s interior.

Figure caption: Deep sea microbial activity under varying pressure. (a) In situ bulk leucine incorporation rates normalized to rates obtained at atmospheric pressure conditions. (b) A microscopic view of a 2000 m sample collected in the Atlantic and incubated under atmospheric pressure conditions. The black halos around the cells are silver grains corresponding to their activities. The highly active cells (indicated by arrows) were rarely found in in situ pressure incubations. (c) Depth-related changes in the metaproteome of three abundant deep sea bacterial taxa (Alteromonas, Bacteroidetes, and SAR202). The number indicates shared and unique up- and down-regulated proteins in different depth zones.

Authors
Chie Amano (University of Vienna, Austria)
Zihao Zhao (University of Vienna, Austria)
Eva Sintes (University of Vienna, IEO-CSIC, Spain)
Thomas Reinthaler (University of Vienna, Austria)
Julia Stefanschitz (University of Vienna, Austria)
Murat Kisadur (University of Vienna, Austria)
Motoo Utsumi (University of Tsukuba, Japan)
Gerhard J. Herndl (University of Vienna, Netherlands Institute for Sea Research)

Twitter @microbialoceanW

How does the competition between phytoplankton and bacteria for iron alter ocean biogeochemical cycles?

Posted by mmaheigan

· Friday, August 26th, 2022

Free-living bacteria play a key role in cycling essential biogeochemical resources in the ocean, including iron, via their uptake, transformation, and release of organic matter throughout the water column. Bacteria process half of the ocean’s primary production, remineralize dissolved organic matter, and re-direct otherwise lost organic matter to higher trophic levels. For these reasons, it is crucial to understand what factors limit the growth of bacteria and how bacteria activities impact global ocean biogeochemical cycles.

In a recent study, Pham and colleagues used a global ocean ecosystem model to dive into how iron limits the growth of free-living marine bacteria, how bacteria modulate ocean iron cycling, and the consequences to marine ecosystems of the competition between bacteria and phytoplankton for iron.

Figure 1: (a) Iron limitation status of bacteria in December, January, and February (DJF) in the surface ocean. Low values (in blue color = close to zero) mean that iron is the limiting factor for the growth of bacteria; (b) Bacterial iron consumption in the upper 120m of the ocean and (c) Changes (anomalies) in export carbon production when bacteria have a high requirement for iron.

Through a series of computer simulations performed in the global ocean ecosystem model, the authors found that iron is a limiting factor for bacterial growth in iron-limited regions in the Southern Ocean, the tropical, and the subarctic Pacific due to the high iron requirement and iron uptake capability of bacteria. Bacteria act as an iron sink in the upper ocean due to their significant iron consumption, a rate comparable to phytoplankton. The competition between bacteria and phytoplankton for iron alters phytoplankton bloom dynamics, ocean carbon export, and the availability of dissolved organic carbon needed for bacterial growth. These results suggest that earth system models that omit bacteria ignore an important organism modulating biogeochemical responses of the ocean to future changes.

Authors:
Anh Le-Duy Pham (Laboratoire d’Océanographie et de Climatologie: Expérimentation et Approches Numériques (LOCEAN), IPSL, CNRS/UPMC/IRD/MNHN, Paris, France)
Olivier Aumont (Laboratoire d’Océanographie et de Climatologie: Expérimentation et Approches Numériques (LOCEAN), IPSL, CNRS/UPMC/IRD/MNHN, Paris, France)
Lavenia Ratnarajah (University of Liverpool, United Kingdom)
Alessandro Tagliabue (University of Liverpool, United Kingdom)

Contrasting N2O fluxes of source vs. sink in western Arctic Ocean during summer 2017

Posted by mmaheigan

· Wednesday, October 20th, 2021

During the western Arctic summer season both physical and biogeochemical features differ with latitude between the Bering Strait and Chukchi Borderland. The southern region (Bering Strait to the Chukchi Shelf) is relatively warm, saline, and eutrophic, due to the intrusion of Pacific waters that bring heat and nutrients in to the western Arctic Ocean (WAO). Because of the Pacific influence, the WAO is one of the most productive stretches of ocean in the world. In contrast, the northern region (Chukchi Borderland to the Canada Basin) is primarily influenced by freshwater originating from sea ice melt and rivers, and is relatively cold, fresh, and oligotrophic. A frontal zone exists between the southern region and northern region (~73°N) due to the distinct physicochemical contrast between mixing Pacific waters and freshwater. These regions support distinct bacterial communities also, making the environmental variations drivers extremely relevant to nitrous oxide (N₂O) dynamics.

A recent study published in Scientific Reports examined the role of the WAO as a source and a sink of atmospheric N₂O. There are obvious differences in N₂O fluxes between southern Chukchi Sea (SC) and northern Chukchi Sea (NC). In the SC (Pacific water characteristics dominate) N₂O emissions act as a net source to the atmosphere (Figure 1a). In the NC (freshwater dominant) absorption of atmospheric N₂O into the water column suggests that this region acts as a net sink (Figure 1a). The positive fluxes of SC occurred with relatively high sea surface temperature (SST), sea surface salinity (SSS), and biogeochemically-derived N₂O production, whereas the negative fluxes of NC were associated with relatively low SST, SSS, and little N₂O production. These linear relationships between N₂O fluxes and environmental variables suggest that summer WAO N₂O fluxes are remarkably sensitive to environmental changes.

Figure 1. (a) Map of the sampling stations using the Ice Breaking R/V Araon during August 2017. The sampling locations were coloured with N2O fluxes (blue to red gradient, see color bar; sink, air → sea (−), and source, sea → air (+). The southern Chukchi Sea (SC) extends from Bering Strait to Chukchi Shelf and the northern Chukchi Sea (NC) extends from Chukchi Borderland and Canada Basin. The frontal zone arises between SC and NC (black dotted line). (b) Illustration showing future changes in the distribution of the WAO N2O flux constrained by the positive feedback scenario of increasing inflow of Pacific waters and rapidly declining sea-ice extent under accelerating Arctic warming.

This study suggests a potential scenario for future WAO changes in terms of accelerating Arctic change. Increasing inflow of the Pacific waters and rapidly declining sea-ice extent are critical. The increasing inflow of warm nutrient-enriched Pacific waters will likely extend the SC N₂O source region northward, increasing productivity, and thereby intensifying nitrification. All of which would lead to a strengthening of the WAO’s role as an N₂O source. A rapid loss of the sea ice extent could ultimately lead to a sea-ice-free NC, and again, a northward shift, which would result in a diminished role of the NC as an N₂O sink (Figure 1b). While improving our understanding of WAO N₂O dynamics, this study suggests both a direction for future work and a clear need for a longer-term study to answer questions about both seasonal variations in these dynamics and possible interannual to climatological trends.

Authors:
Jang-Mu Heo (Department of Marine Science, Incheon National University)
Sang-Min Eom (Department of Marine Science, Incheon National University)
Alison M. Macdonald (Woods Hole Oceanographic Institution)
Hyo-Ryeon Kim (Department of Marine Science, Incheon National University)
Joo-Eun Yoon (Department of Marine Science, Incheon National University)
Il-Nam Kim (Department of Marine Science, Incheon National University)

Partitioning carbon export into particulate and dissolved pools from biogeochemical profiling float observations

Posted by mmaheigan

· Thursday, December 17th, 2020

Carbon export from the surface into the deep ocean via the biological pump is a significant sink for atmospheric carbon dioxide. The relative contributions of sinking particles—particulate organic carbon (POC) and dissolved organic carbon (DOC)—to the total export affect the efficiency of carbon export.

In a recent study published in Global Biogeochemical Cycles, the authors used measurements from biogeochemical profiling floats in the Northeast Pacific from 2009 to 2017 to estimate net community production (NCP), an analog for carbon export. In order to close three tracer budgets (nitrate, dissolved inorganic carbon, and total alkalinity), the authors combined these float measurements with data from the Ocean Station Papa mooring and recently developed algorithms for carbonate system parameters. By constraining end-member nutrient ratios of the POC and DOC produced, this multi-tracer approach was used to estimate regional NCP across multiple depth horizons throughout the annual cycle, partition NCP into the POC and DOC contributions, and calculate particulate inorganic carbon (PIC) production, a known ballast material for sinking particles (Figure 1). The authors also estimated POC attenuation with depth, POC export across deeper horizons, and in situ export efficiency via a particle backscatter-based approach.

With the advent of “fully-loaded” biogeochemical profiling floats equipped with nitrate, oxygen, pH and bio-optical sensors, this approach may be used to assess the magnitude and efficiency of carbon export in other ocean regions from a single platform, which will greatly reduce the risks and costs associated with traditional ship-based measurements, while broadening the spatiotemporal scales of observation.

Figure caption: Climatological mean NCP (blue line) over the entire study period (2009-2017); the POC portion of NCP (filled blue area), the DOC portion (white space) and PIC production rate (red line), in the mixed layer (left), and the euphotic zone (right). The numbers in parentheses are the integrated annual NCP rates for each curve and uncertainty reported was determined using a Monte Carlo approach.

Authors:
William Haskell (MBARI, now Mote Marine Laboratory)
Andrea Fassbender (MBARI, now PMEL)
Jacki Long (MBARI)
Joshua Plant (MBARI)

Modern OMZ copepod dynamics provide analog for future oceans

Posted by mmaheigan

· Thursday, July 23rd, 2020

Global warming increases ocean deoxygenation and expands the oxygen minimum zone (OMZ), which has implications for major zooplankton groups like copepods. Reduced oxygen levels may impact individual copepod species abundance, vertical distribution, and life history strategy, which is likely to perturb intricate oceanic food webs and export processes. In a study recently published in Biogeosciences, authors conducted vertically-stratified day and night MOCNESS tows (0-1000 m) during four cruises (2007-2017) in the Eastern Tropical North Pacific, sampling hydrography and copepod distributions in four locations with different water column oxygen profiles and OMZ intensity (i.e. lowest oxygen concentration and its vertical extent in a profile). Each copepod species exhibited a different vertical distribution strategy and physiology associated with oxygen profile variability. The study identified sets of species that (1) changed their vertical distributions and maximum abundance depth associated with the depth and intensity of the OMZ and its oxycline inflection points, (2) shifted their diapause depth, (3) adjusted their diel vertical migration, especially the nighttime upper depth, or (4) expanded or contracted their depth range within the mixed layer and upper part of the thermocline in association with the thickness of the aerobic epipelagic zone (habitat compression concept) (Figure 1). Distribution depths for some species shifted by 10’s to 100’s of meters in different situations, which also had metabolic (and carbon flow) implications because temperature decreased with depth. This observed present-day variability may provide an important window into how future marine ecosystems will respond to deoxygenation.

Figure caption: Schematic diagram showing how future OMZ expansion may affect zooplankton distributions, based on present-day responses to OMZ variability. The dashed line indicates diel vertical migration (DVM) and highlights the shoaling of the nighttime depth as the aerobic habitat is compressed. The lower oxycline community and the diapause layer for some species, associated with a specific oxygen concentration, may deepen as the OMZ expands.

Authors:
Karen F. Wishner (University of Rhode Island)
Brad Seibel (University of South Florida)
Dawn Outram (University of Rhode Island)

Surface bacterial communities respond to rapid changes in the western Arctic

Posted by mmaheigan

· Tuesday, January 7th, 2020

During the western Arctic summer open water season, latitudinal differences in the physical and biogeochemical features of the surface water are apparent from the Bering Strait to the Chukchi Borderland. Lower latitude regions (i.e. Bering Strait to Chukchi Shelf) are primarily driven by the inflow of Pacific waters that supply nutrients and heat, leading to high primary production. Conversely, the higher latitude regions (i.e. Chukchi Borderland and Canada Basin) are relatively cold, fresh, and oligotrophic because the surface layer is influenced by freshwater inputs from melting ice and rivers via the Beaufort Gyre. Mixing of the two surface water masses in the western Arctic produces a physicochemical frontal zone (FZ) in the Chukchi Sea.

In a recent study published in Scientific Reports, authors used observations from summer 2017 to investigate latitudinal variations in bacterial community composition in surface waters between the Bering Strait and Chukchi Borderland and the underlying processes driving the changes. Results indicate three distinctive communities: 1) Southern Chukchi (SC) bacterial communities are associated with nutrient-rich conditions, including genera such as Sulfitobacter; 2) a northern Chukchi (NC) bacterial community that dominated by SAR clades, Flavobacterium, Paraglaciecola, and Polaribacter, genera associated with low nutrients and sea ice conditions. If climate-driven changes in the western Arctic continue along the same trajectory, it’s likely we will see altered bacterial communities. If the impact of warm, nutrient-rich Pacific water inflows dominates, it is likely that the productive SC region will expand and the FZ will move northward, leading to nutrient enrichment in the western Arctic (Figure 1). In response, bacterial communities would be dominated by organic matter decomposers, such as Sulfitobacter, due to high primary productivity. However, if the impact of sea-ice meltwater dominates, then the oligotrophic NC region will expand and the FZ will move southward, leading to nutrient depletion in western Arctic surface waters (Figure 1). Continued monitoring in this region will enhance our understanding of how bacterial communities respond (Figure 1b) to a rapidly changing western Arctic Ocean.

Figure 1. (a) Map of the August 2017 Ice Breaker RV Araon western Arctic Ocean sampling stations used in this study. The basemap shows the Chl-a concentration contour (blue to red background colors). Pink, green, and blue circles represent stations in the South Chukchi (SC), Frontal Zone (FZ), and Northern Chukchi (NC) regions. (b) Schematic diagram of surface bacterial community distribution in response to future western Arctic Ocean changes.

Authors:
Il-Nam Kim (Department of Marine Science, Incheon National University)
Sung-Ho Kang (Korea Polar Research Institute)
Eun Jin Yang (Korea Polar Research Institute)

The Equatorial Undercurrent influences the fate of the Oxygen Minimum Zone in the Pacific

Posted by mmaheigan

· Tuesday, November 12th, 2019

While the ocean as a whole is losing oxygen due to warming, oxygen minimum zones (OMZs) are maintained by a delicate balance of biological and physical processes; it is unclear how each one of them is going to evolve in the future. Changes to OMZs could affect the global uptake of carbon, the generation of greenhouse gases, and interactions among marine life. Current generation coarse-resolution (~1°) climate models compromise the ability to simulate low-oxygen waters and their response to climate change in the future because they fail to reproduce a major ocean current, the Equatorial Undercurrent (EUC). These shortcomings lead to an overly tilted upper oxygen minimum zone (OMZ) (Figure 1), thus exaggerating sensitivity to circulation changes and overwhelming other key processes like diffusion and biology. The EUC also plays a vital role in feeding the eastern Pacific upwelling region, connecting it to global climate variability.

Figure: Top: The boundary of the Oxygen Minimum Zone (OMZ) along the Equator is unrealistically tilted for current generation (coarse resolution) climate models, and improves with increased horizontal resolution. The tilt is due to a bad representation of the Equatorial Undercurrent in the coarse model, also seen in other coarse models. The exaggerated tilt of the OMZ boundary at the Equator leads to increased inter-annual variability of the depth of the upper OMZ boundary, via changes in the zonal flow (left). This phenomenon is found in most CMIP5 models (right) and could be responsible for the current inability to predict the change in OMZ extent for the next century.

A recent high‐resolution climate model study in Geophysical Research Letters significantly improved the representation of both the EUC and OMZ, suggesting that the EUC is a key player in OMZ variability. This study emphasizes the importance of improving transport processes in global circulation models to better simulate oxygen distribution and predict future OMZ extent. The results of this study imply that the fundamental dynamics maintaining this key ocean current could be categorically misrepresented in the current generation of climate models, potentially influencing the ability to predict future climate variability and trends.

Authors:
Julius J.M. Busecke (Princeton University)
Laure Resplandy (Princeton University)
John P. Dunne (NOAA/GFDL)

Biogeochemical controls of surface ocean phosphate

Posted by mmaheigan

· Tuesday, November 12th, 2019

Phosphorus availability is important for phytoplankton growth and more broadly ocean biogeochemical cycles. However, phosphate concentration is often below the analytical detection limit of the standard auto-analyzer technique. Thus, we know little about geographic phosphate variation across most low latitude regions. To address this issue, a global collaboration of scientists conducted a study published in Science Advances on combined phosphate measurements using high-sensitivity methods that yielded a detailed map of surface phosphate (Figure 1).

Figure 1: Fine-scale global variation of surface phosphate. Surface phosphate measured using high-sensitivity techniques revealed previously unrecognized low latitude differences in phosphate drawdown.

The study’s new globally expansive phosphate data set revealed previously unrecognized low-phosphate areas, including large regions of the Pacific Ocean—really low phosphate in the western North Pacific and to a lesser extent in the South Pacific. Although atmospheric iron input and nitrogen fixation are commonly described as regulators of surface phosphate, this study shows that shifts in the elemental stoichiometry (N:P:Fe) of the vertical nutrient supply play an additional role. Previous studies and climate models have suggested that the availability of phosphate is a first-order driver of ocean biogeochemical changes. Interestingly, this study suggests that marine ecosystems are more resilient to phosphate stress than previously thought. These findings underscore the importance of accurately quantifying nutrients at low concentrations for understanding the regulation of ocean ecosystem processes and biogeochemistry now and under future climate conditions.

And the data are of course available in BCO-DMO!

Authors:
Adam C. Martiny (University of California, Irvine)
Michael W. Lomas (Bigelow Laboratory for Ocean Sciences)
Weiwei Fu (University of California, Irvine)
Philip W. Boyd (University of Tasmania)
Yuh-ling L. Chen (National Sun Yat-sen University)
Gregory A. Cutter (Old Dominion University)
Michael J. Ellwood (Australian National University)
Ken Furuya (The University of Tokyo)
Fuminori Hashihama (Tokyo University of Marine Science and Technology)
Jota Kanda (Tokyo University of Marine Science and Technology)
David M. Karl (University of Hawaii)
Taketoshi Kodama (Japan Fisheries Research and Education Agency)
Qian P. Li (Chinese Academy of Sciences)
Jian Ma (Xiamen University)
Thierry Moutin (Université de Toulon)
E. Malcolm S. Woodward (Plymouth Marine Laboratory)
J. Keith Moore (University of California, Irvine)

The arsenic respiratory cycle in pelagic waters of Oxygen Deficient Zones

Posted by mmaheigan

· Wednesday, October 30th, 2019

Oxygen Deficient Zones (ODZs) are naturally occurring functionally anoxic regions of the open ocean which can act as proxies for early Earth’s anoxic ocean. Without free oxygen, microorganisms in these regions use alternative electron acceptors to oxidize organic material. These functionally anoxic regions are also hotspots for chemoautotrophic pathways. Some microorganisms can use arsenic based compounds to oxidize organic material, and others can couple nitrate reduction with arsenic oxidation supporting autotrophic carbon fixation thus linking arsenic respiration with carbon and nitrogen cycling. While arsenic concentrations in modern oceans are relatively low, the Precambrian ocean likely had periods of high arsenic concentrations. Integrating over time and space of anoxic waters, arsenic-based metabolisms may have had significant implications for the biogeochemical cycling of not only arsenic, but also carbon and nitrogen.

Figure 1: Arsenotrophic genes identified in the Eastern Tropical North Pacific Oxygen Deficient Zone. (A) Genomic complement for dissimilatory arsenate reduction assembled from metagenomes which likely supports respiration of organic matter. (B) Genomic complement for putative chemoautotrophic arsenite oxidation pathway assembled from metagenomes which may couple with nitrate reduction to support organic matter production. (C) Relative abundance of genes associated with arsenite oxidase (aioA), dissimilatory arsenate reduction (arrA), and forward dissimilatory sulfite reductase (dsrA) associated with sulfur reduction; abundance shown as a relative contribution to the total microbial community as estimated by abundance of RNA polymerase genes (rpoB). The genes arrA and forward-dsrA are more abundant in the particulate fraction, whereas aioA is more abundant in the free-living fraction. (D) Relative abundance of genes in the microbial community for the more abundant genes aioA-like and reverse form of dsrA associated with sulfur oxidation. aioA-like genes are relatively more abundant within the particulate fraction, with no strong partitioning between fractions identified for the reverse-dsrA genes. Arsenical reduction and chemoautotrophic arsenical oxidation are likely performed by different microbial groups within the ODZ communities.

Recent work in PNAS identified gene sequences for a complete arsenic respiratory cycle from Eastern Tropical North Pacific (ETNP) ODZ metagenomes. The authors identified arsenotrophic genes for dissimilatory arsenate reduction from one group of microorganisms and genes for a putative chemoautotrophic arsenite oxidation pathway from another group within the ETNP ODZ microbial community. Analysis of genomic sequences from a free-living sample and from particulate-associated sample indicate niche differentiation of these pathways—arsenate reduction genes enriched within the particulate fraction and arenite oxidation enriched in the free-living water column. In addition to the presence of these genes in metagenomes, the authors identified the active expression of these arsenotrophic genes in publicly available metatranscriptomes from the ETNP and Eastern Tropical South Pacific ODZs. Theyalso found an abundance of sequences in the ETNP ODZ for the gene aioA-like, which is a closely related enzyme to arsenite oxidase (aioA), but with an unconfirmed function. The identification of these actively expressed genes in modern ODZs enables further investigation of these cycles that were likely important in early oceans. These findings also highlight that there are still yet to be discovered respiratory pathways in ODZs. Arsenotrophy, in conjunction with other niche respiratory pathways – both known and as yet undiscovered – likely sum to a considerable contribution of energy flow and elemental cycling through these anoxic systems.

Authors:
Jaclyn Saunders (University of Washington; present affiliation Woods Hole Oceanographic Institution)
Clara Fuchsman (University of Washington; present affiliation Horn Point Laboratory)
Cedar McKay & Gabrielle Rocap (University of Washington)

See related University of Washington press-release

Phytoplankton bloom from molten lava

Posted by mmaheigan

· Wednesday, September 18th, 2019

During June-August 2018, the oligotrophic North Pacific Ocean received an enormous quantity of nutrients in the form of molten lava, delivered by the erupting Kilauea on the big island of Hawaii. A phytoplankton bloom formed in response to the input of lava and an expedition was rapidly mobilized to determine its composition and the relevant biogeochemistry. We found that in addition to the nutrients derived from lava, exogenous nitrate was also present in the surface waters. Remotely operated vehicle observations in September 2019 by scientists at the Woods Hole Oceanographic Institution showed that lava from the 2018 eruption had reached depths of 700 m. Therefore, enabled by the intensity of the eruption and the island’s steep bathymetry, lava flows were able to extend below the thermocline and penetrate into nitrate-rich waters. Based on isotopic signatures of nitrate in the bloom, we inferred that heating of deep ocean waters resulted in the formation of buoyant seawater plumes, which rose to the sea surface. The rapid response expedition in July 2018 provided a unique opportunity to see first-hand how a massive input of exogenous nutrients alters marine ecosystems attuned to oligotrophic conditions.

Read more:
Ducklow, H. and T. Plank (06 Sep 2019) Volcano-stimulated marine photosynthesis. Science. Vol. 365, Issue 6457, pp. 978-979
DOI: 10.1126/science.aay8088>

Wilson, S. et al. (06 Sep 2019) Kīlauea lava fuels phytoplankton bloom in the North Pacific Ocean. Science Vol. 365, Issue 6457, pp.1040-1044
DOI: 10.1126/science.aax4767

Funding for the Ocean Carbon & Biogeochemistry Project Office is provided by the National Science Foundation (NSF) and the National Aeronautics and Space Administration (NASA). The OCB Project Office is housed at the Woods Hole Oceanographic Institution.

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