Understanding the unique biogeochemistry of the Mediterranean Sea: Insights from a coupled phosphorus and nitrogen model

The Mediterranean Sea (MS) is an oligotrophic basin whose offshore water column exhibits low dissolved inorganic phosphorus (P) and nitrogen (N) concentrations, unusually high nitrate (NO3) to phosphate (PO4) ratios, and distinct biogeochemical differences between the Western Mediterranean Sea (WMS) and Eastern Mediterranean Sea (EMS). A new mass balance model of P and N cycling in the WMS is coupled to a pre‐existing EMS model to understand these biogeochemical features. Estimated land‐derived inputs of reactive P and N to the WMS and EMS are similar per unit surface area, but marine inputs are 4 to 5 times greater for the WMS, which helps explain the approximately 3 times higher primary productivity of the WMS. The lateral inputs of marine sourced inorganic and organic P support significant fractions of new production in the WMS and EMS, similar to subtropical gyres. The mass balance calculations imply that the MS is net heterotrophic: dissolved organic P and N entering the WMS and EMS, primarily via the Straits of Gibraltar and Sicily, are mineralized to PO4 and NO3 and subsequently exported out of the basin by the prevailing anti‐estuarine circulation. The high deepwater (DW) molar NO3:PO4 ratios reflect the high reactive N:P ratio of inputs to the WMS and EMS, combined with low denitrification rates. The lower DW NO3:PO4 ratio of the WMS (21) compared to the EMS (28) reflects lower reactive N:P ratios of inputs to the WMS, including the relatively low N:P ratio of Atlantic surface water flowing into the WMS

Gastrointestinal-projecting neurones in the dorsal motor nucleus of the vagus exhibit direct and viscerotopically organized sensitivity to orexin

Peer Reviewedhttp://deepblue.lib.umich.edu/bitstream/2027.42/65136/1/jphysiol.2002.029546.pd

Modelling terrigenous DOC across the north west European Shelf: Fate of riverine input and impact on air-sea CO2 fluxes

Terrigenous carbon in aquatic systems is increasingly recognised as an important part of the global carbon cycle. Despite this, the fate and distribution of terrigenous dissolved organic carbon (tDOC) in coastal and oceanic systems is poorly understood. We have implemented a theoretical framework for the degradation of tDOC across the land to ocean continuum in a 3D hydrodynamical-biogeochemical model on the North West European Shelf. A key feature of this model is that both photochemical and bacterial tDOC degradation rates are age dependant constituting an advance in our ability to describe carbon cycling in the marine environment. Over the time period 1986-2015, 182±17 Gmol yr− 1 of riverine tDOC is input to the shelf. Results indicate that bacterial degradation is by far the most important process in removing tDOC on the shelf, contributing to 73±6 % (132±11 Gmol yr− 1 ) of the total removal flux, while 21±3 % (39±6 Gmol yr− 1 ) of riverine tDOC was advected away from the shelf and photochemical degradation removing 5±0.5 % of the riverine flux. Explicitly including tDOC in the model decreased the air-sea carbon dioxide (CO2) flux by 112±8 Gmol yr− 1 (4±0.4 %), an amount approximately equivalent to the CO2 released by the UK chemical industry in 2020. The reduction is equivalent to 62 % of the riverine tDOC input to the shelf while approximately 17 % of riverine input is incorporated into the foodweb. This work can improve the assumptions of the fate of tDOC by Earth System Models and demonstrates that the inclusion of tDOC in models can impact ecosystem dynamics and change predicted global carbon budgets for the ocean

Deep-Ocean dissolved organic matter reactivity along the Mediterranea Sea: does size matter?

Original research paperDespite of the major role ascribed to marine dissolved organic matter (DOM) in the global carbon cycle, the reactivity of this pool in the dark ocean is still poorly understood. Present hypotheses, posed within the size-reactivity continuum (SRC) and the microbial carbon pump (MCP) conceptual frameworks, need further empirical support. Here, we provide field evidence of the soundness of the SRC model. We sampled the high salinity core-of-flow of the Levantine Intermediate Water along its westward route through the entire Mediterranean Sea. At selected sites, DOM was size-fractionated in apparent high (aHMW) and low (aLMW) molecular weight fractions using an efficient ultrafiltration cell. A percentage decline of the aHMW DOM from 68–76% to 40–55% was observed from the Levantine Sea to the Strait of Gibraltar in parallel with increasing apparent oxygen utilization (AOU). DOM mineralization accounted for 30±3% of the AOU, being the aHMW fraction solely responsible for this consumption, verifying the SRC model in the field. We also demonstrate that, in parallel to this aHMW DOM consumption, fluorescent humic-like substances accumulate in both fractions and protein-like substances decline in the aLMW fraction, thus indicating that not only size matters and providing field support to the MCP modelHOTMIX (grant number CTM2011–30010-C02 01-MAR and 02-MAR) and the project FERMIO (MINECO, CTM2014-57334-JIN), both co-financed with FEDER funds; (reference BES-2012- 056175) from the Spanish Ministry of Economy, Industry and Competitivenes; the project MODMED from CSIC (PIE, 201730E020) and CSIC Program “Junta para la Ampliación de Estudios” co-financed by the ESF (reference JAE DOC 040)Versión del editor2,92

Extrinsic primary afferent signalling in the gut

Visceral sensory neurons activate reflex pathways that control gut function and also give rise to important sensations, such as fullness, bloating, nausea, discomfort, urgency and pain. Sensory neurons are organised into three distinct anatomical pathways to the central nervous system (vagal, thoracolumbar and lumbosacral). Although remarkable progress has been made in characterizing the roles of many ion channels, receptors and second messengers in visceral sensory neurons, the basic aim of understanding how many classes there are, and how they differ, has proven difficult to achieve. We suggest that just five structurally distinct types of sensory endings are present in the gut wall that account for essentially all of the primary afferent neurons in the three pathways. Each of these five major structural types of endings seems to show distinctive combinations of physiological responses. These types are: 'intraganglionic laminar' endings in myenteric ganglia; 'mucosal' endings located in the subepithelial layer; 'muscular–mucosal' afferents, with mechanosensitive endings close to the muscularis mucosae; 'intramuscular' endings, with endings within the smooth muscle layers; and 'vascular' afferents, with sensitive endings primarily on blood vessels. 'Silent' afferents might be a subset of inexcitable 'vascular' afferents, which can be switched on by inflammatory mediators. Extrinsic sensory neurons comprise an attractive focus for targeted therapeutic intervention in a range of gastrointestinal disorders.Australian National Health and Medical Research Counci

A biogeochemical model for phosphorus and nitrogen cycling in the Eastern Mediterranean Sea: Part 2: Response of nutrient cycles and primary production to anthropogenic forcing: 1950-2000.

Anthropogenic inputs of nutrient phosphorus (P) and nitrogen (N) to the Eastern Mediterranean Sea (EMS) increased significantly after 1950. Nonetheless, the EMS remained ultra-oligotrophic, with eutrophication only affecting a restricted number of nearshore areas. To better understand this apparent contradiction, we reconstructed the external inputs of reactive P and N to the EMS for the period 1950 to 2000. Although the inputs associated with atmospheric deposition and river discharge more than doubled, the inflow of surface water from the Western Mediterranean Sea (WMS) remained the dominant source of nutrient P and N to the EMS during the second half of the 20th century. The combined external input of reactive P rose by 24% from 1950 to 1985, followed by a slight decline. In contrast, the external reactive N input increased continuously from 1950 to 2000, with a 62% higher input in 2000 compared to 1950. When imposing the reconstructed inputs to the dynamic model of P and N cycling in the EMS developed in the companion paper, a maximum increase of primary production of only 16% is predicted. According to the model, integrated over the period 1950-2000, outflow of Levantine Intermediate Water (LIW) to the WMS exported the equivalent of about one third of the P supplied in excess of the 1950 input, while another one third was translocated to the Eastern Mediterranean Deep Water (EMDW). Together, both mechanisms efficiently counteracted enhanced P input to the EMS, by drawing nutrient P away from primary producers in the surface waters. Furthermore, between 1950 and 2000, inorganic and organic dissolved N:P ratios increased in all water masses. Thus, the EMS became even more P limited because of anthropogenic nutrient inputs. A model simulation incorporating the circulation changes accompanying the Eastern Mediterranean Transient (EMT) between 1987 and 2000 yielded a 4% increase of EMS primary productivity relative to the baseline scenario

A biogeochemical model for phosphorus and nitrogen cycling in the Eastern Mediterranean Sea: Part 1: Model development, initialization and sensitivity

The Eastern Mediterranean Sea (EMS) is the largest marine basin whose annual primary productivity is limited by phosphorus (P) rather than nitrogen (N). The basin is nearly entirely land-locked and receives substantial external nutrient fluxes, comparable for instance to those of the Baltic Sea. The biological productivity of the EMS, however, is among the lowest observed in the oceans. The water column exhibits very low P and N concentrations with N:P ratios in excess of the Redfield value. These unique biogeochemical features are analyzed using a mass balance model of the coupled P and N cycles in the EMS. The present paper describes the conceptual basis, quantitative implementation and sensitivity of the model. The model is initialized for the year 1950, that is, prior to the large increase in anthropogenic nutrient loading experienced by the EMS during the second half of the 20th century. In the companion paper, the model is used to simulate the P and N cycles during the period 1950-2000. The 1950 model set-up and sensitivity analyses support the following conclusions.(1)Phosphorus-limited primary production in the EMS is most sensitive to the P exchanges with the Western Mediterranean Sea (WMS) associated with the anti-estuarine circulation of the EMS. The supply of P through the Straits of Sicily is mainly under the form of dissolved organic P (DOP), while dissolved inorganic P (PO4) is primarily exported to the WMS. The efficient export of PO4 to the WMS maintains the EMS in its ultra-oligotrophic state.(2)Inorganic molar N:P ratios in excess of the 16:1 Redfield value observed in the water column reflect higher-than-Redfield N:P ratios of the external inputs, combined with negligible denitrification. Model simulations imply that the denitrification flux would have to increase by at least a factor of 14, relative to the 1950 flux, in order for the inorganic N:P ratio of the deep waters to approach the Redfield value.(3)The higher-than-Redfield N:P ratios of dissolved and particulate organic matter in the EMS further imply the preferential regeneration of P relative to N during organic matter decomposition

Neuroanatomy of extrinsic afferents supplying the gastrointestinal tract

Here we discuss the neuroanatomy of extrinsic gastrointestinal (GI) afferent neurones, the relationship between structure and function and the role of afferents in disease. Three pathways connect the gut to the central nervous system: vagal afferents signal mainly from upper GI regions, pelvic afferents mainly from the colorectal region and splanchnic afferents from throughout. Vagal afferents mediate reflex regulation of gut function and behaviour, operating mainly at physiological levels. There are two major functional classes - tension receptors, responding to muscular contraction and distension, and mucosal receptors. The function of vagal endings correlates well with their anatomy: tracing studies show intramuscular arrays (IMAs) and intraganglionic laminar endings (IGLEs); IGLEs are now known to respond to tension. Functional mucosal receptors correlate with endings traced to the lamina propria. Pelvic afferents serve similar functions to vagal afferents, and additionally mediate both innocuous and noxious sensations. Splanchnic afferents comprise mucosal and stretch-sensitive afferents with low thresholds in addition to high-threshold serosal/mesenteric afferents suggesting diverse roles. IGLEs, probably of pelvic origin, have been identified recently in the rectum and respond similarly to gastric vagal IGLEs. Gastrointestinal afferents may be sensitized or inhibited by chemical mediators released from several cell types. Whether functional changes have anatomical correlates is not known, but it is likely that they underlie diseases involving visceral hypersensitivity.Berthoud, H R ; Blackshaw, L A ; Brookes, S J H ; Grundy,