Indicators of sustainability of agriculture and livestock in Spain - Indicadores de sostenibilidad de la agricultura y ganadería española

Sustainability is an adjective used to characterize agriculture according to the degree of fulfillment of goals. Those goals are related to agro-ecological, environmental and socio-economic dimensions. Sustainability is a dynamic and temporal character. In absolute terms there is not an ending value because it changes as its dimensions make it. Spain is one of the main agricultural countries of the European Union both in terms of crop land and value of productions. The object of this study is to present a methodology of sustainability account to be incorporated into national statistical and to assess their performance in the course of the years. For that reason the data sources used have been the statistics of the Department of Agriculture and from others database. We presented a set of indicators of sustainability and its evaluation in a time series of at least 30 years. The trend analysis offers the evolution of the numerical values of the indicators in terms of efficiency, physical units used for a unit of product or its value in euros. The analyzed crops have been: wheat, barley, maize, sunflower, sugar beet, wine grape, olive oil, citrus, melon and tomato. Physical indicators were: land, water, energy, erosion, soil organic matter, and carbon balance; socio-economic indicators were: agricultural final production, prices, income, employment and use of fertilizers. In general, all crops increased their productive efficiency, higher in irrigated than on dry land. Spanish agricultural carbon sequestration capacity has multiplied by five in the last seventy years, as a result of the increase in the productivity of crops, in terms of total biomass and the modification of the soil management techniques. Livestock sector presents data of pork, broilers and laying hen. Those showed an improvement in efficiency and economic indicators. Overall we can say that Spanish agriculture and livestock subsector have a tendency towards sustainability, being its main threats extreme meteorological factors and the instability of todays markets.

Functional response of Ulmus minor Mill. to drought, flooding and dutch elm disease

Ulmus minor es una especie arbórea originaria de Europa cuyas poblaciones han sido diezmadas por el hongo patógeno causante de la enfermedad de la grafiosis. La conservación de los olmos exige plantearse su propagación a través de plantaciones y conocer mejor su ecología y biología. Ulmus minor es un árbol de ribera, pero frecuentemente se encuentra alejado del cauce de arroyos y ríos, donde la capa freática sufre fuertes oscilaciones. Por ello, nuestra hipótesis general es que esta especie es moderadamente resistente tanto a la inundación como a la sequía. El principal objetivo de esta tesis doctoral es entender desde un punto de vista funcional la respuesta de U. minor a la inundación, la sequía y la infección por O. novo-ulmi; los factores que posiblemente más influyen en la distribución actual de U. minor. Con este objetivo se persigue dar continuidad a los esfuerzos de conservación de esta especie que desde hace años se dedican en varios centros de investigación a nivel mundial, ya que, entender mejor los mecanismos que contribuyen a la resistencia de U. minor ante la inoculación con O. novo-ulmi y factores de estrés abiótico ayudará en la selección y propagación de genotipos resistentes a la grafiosis. Se han planteado tres experimentos en este sentido. Primero, se ha comparado la tolerancia de brinzales de U. minor y U. laevis – otro olmo ibérico – a una inmersión controlada con el fin de evaluar su tolerancia a la inundación y comprender los mecanismos de aclimatación. Segundo, se ha comparado la tolerancia de brinzales de U. minor y Quercus ilex – una especie típica de ambientes Mediterránea secos – a la falta de agua en el suelo con el fin de evaluar el grado de tolerancia y los mecanismos de aclimatación a la sequía. El hecho de comparar dos especies contrastadas responde al interés en entender mejor cuales son los procesos que conducen a la muerte de una planta en condiciones de sequía – asunto sobre el que hay una interesante discusión desde hace algunos años. En tercer lugar, con el fin de entender mejor la resistencia de algunos genotipos de U. minor a la grafiosis, se han estudiado las diferencias fisiológicas y químicas constitutivas e inducidas por O. novo-ulmi entre clones de U. minor seleccionados a priori por su variable grado de resistencia a esta enfermedad. En el primer experimento se observó que los brinzales de U. minor sobrevivieron 60 días inmersos en una piscina con agua no estancada hasta una altura de 2-3 cm por encima del cuello de la raíz. A los 60 días, los brinzales de U. laevis se sacaron de la piscina y, a lo largo de las siguientes semanas, fueron capaces de recuperar las funciones fisiológicas que habían sido alteradas anteriormente. La conductividad hidráulica de las raíces y la tasa de asimilación de CO2 neta disminuyeron en ambas especies. Por el contrario, la tasa de respiración de hojas, tallos y raíces aumentó en las primeras semanas de la inundación, posiblemente en relación al aumento de energía necesario para desarrollar mecanismos de aclimatación a la inundación, como la hipertrofia de las lenticelas que se observó en ambas especies. Por ello, el desequilibrio del balance de carbono de la planta podría ser un factor relevante en la mortalidad de las plantas ante inundaciones prolongadas. Las plantas de U. minor (cultivadas en envases de 16 litros a media sombra) sobrevivieron por un prolongado periodo de tiempo en verano sin riego; la mitad de las plantas murieron tras 90 días sin riego. El cierre de los estomas y la pérdida de hojas contribuyeron a ralentizar las pérdidas de agua y tolerar la sequía en U. minor. Las obvias diferencias en tolerancia a la sequía con respecto a Q. ilex se reflejaron en la distinta capacidad para ralentizar la aparición del estrés hídrico tras dejar de regar y para transportar agua en condiciones de elevada tensión en el xilema. Más relevante es que las plantas con evidentes síntomas de decaimiento previo a su muerte exhibieron pérdidas de conductividad hidráulica en las raíces del 80% en ambas especies, mientras que las reservas de carbohidratos apenas variaron y lo hicieron de forma desigual en ambas especies. Árboles de U. minor de 5 y 6 años de edad (plantados en eras con riego mantenido) exhibieron una respuesta a la inoculación con O. novo-ulmi consistente con ensayos previos de resistencia. La conductividad hidráulica del tallo, el potencial hídrico foliar y la tasa de asimilación de CO2 neta disminuyeron significativamente en relación a árboles inoculados con agua, pero solo en los clones susceptibles. Este hecho enlaza con el perfil químico “más defensivo” de los clones resistentes, es decir, con los mayores niveles de suberina, ácidos grasos y compuestos fenólicos en estos clones que en los susceptibles. Ello podría restringir la propagación del hongo en el árbol y preservar el comportamiento fisiológico de los clones resistentes al inocularlos con el patógeno. Los datos indican una respuesta fisiológica común de U. minor a la inundación, la sequía y la infección por O. novo-ulmi: pérdida de conductividad hidráulica, estrés hídrico y pérdida de ganancia neta de carbono. Pese a ello, U. minor desarrolla varios mecanismos que le confieren una capacidad moderada para vivir en suelos temporalmente anegados o secos. Por otro lado, el perfil químico es un factor relevante en la resistencia de ciertos genotipos a la grafiosis. Futuros estudios deberían examinar como este perfil químico y la resistencia a la grafiosis se ven alteradas por el estrés abiótico. ABSTRACT Ulmus minor is a native European elm species whose populations have been decimated by the Dutch elm disease (DED). An active conservation of this species requires large-scale plantations and a better understanding of its biology and ecology. U. minor generally grows close to water channels. However, of the Iberian riparian tree species, U. minor is the one that spread farther away from rivers and streams. For these reasons, we hypothesize that this species is moderately tolerant to both flooding and drought stresses. The main aim of the present PhD thesis is to better understand the functional response of U. minor to the abiotic stresses – flooding and drought – and the biotic stress – DED – that can be most influential on its distribution. The overarching goal is to aid in the conservation of this emblematic species through a better understanding of the mechanisms that contribute to resistance to abiotic and biotic stresses; an information that can help in the selection of resistant genotypes and their expansion in large-scale plantations. To this end, three experiments were set up. First, we compared the tolerance to experimental immersion between seedlings of U. minor and U. laevis – another European riparian elm species – in order to assess their degree of tolerance and understand the mechanisms of acclimation to this stress. Second, we investigated the tolerance to drought of U. minor seedlings in comparison with Quercus ilex (an oak species typical of dry Mediterranean habitats). Besides assessing and understanding U. minor tolerance to drought at the seedling stage, the aim was to shed light into the functional alterations that trigger drought-induced plant mortality – a matter of controversy in the last years. Third, we studied constitutive and induced physiological and biochemical differences among clones of variable DED resistance, before and following inoculation with Ophiostoma novo-ulmi. The goal is to shed light into the factors of DED resistance that is evident in some genotypes of U. minor, but not others. Potted seedlings of U. minor survived for 60 days immersed in a pool with running water to approximately 2-3 cm above the stem collar. By this time, U. minor seedlings died, whereas U. laevis seedlings moved out of the pool were able to recover most physiological functions that had been altered by flooding. For example, root hydraulic conductivity and leaf photosynthetic CO2 uptake decreased in both species; while respiration initially increased with flooding in leaves, stems and roots possibly to respond to energy demands associated to mechanisms of acclimation to soil oxygen deficiency; as example, a remarkable hypertrophy of lenticels was soon observed in flooded seedlings of both species. Therefore, the inability to maintain a positive carbon balance somehow compromises seedling survival under flooding, earlier in U. minor than U. laevis, partly explaining their differential habitats. Potted seedlings of U. minor survived for a remarkable long time without irrigation – half of plants dying only after 90 days of no irrigation in conditions of high vapour pressure deficit typical of summer. Some mechanisms that contributed to tolerate drought were leaf shedding and stomata closure, which reduced water loss and the risk of xylem cavitation. Obviously, U. minor was less tolerant to drought than Q. ilex, differences in drought tolerance resulting mostly from the distinct capacity to postpone water stress and conduct water under high xylem tension among species. More relevant was that plants of both species exhibited similar symptoms of root hydraulic failure (i.e. approximately 80% loss of hydraulic conductivity), but a slight and variable depletion of non-structural carbohydrate reserves preceding dieback. Five- and six-year-old trees of U. minor (planted in the field with supplementary watering) belonging to clones of contrasted susceptibility to DED exhibited a different physiological response to inoculation with O. novo-ulmi. Stem hydraulic conductivity, leaf water potential and photosynthetic CO2 uptake decreased significantly relative to control trees inoculated with water only in DED susceptible clones. This is consistent with the “more defensive” chemical profile observed in resistant clones, i.e. with higher levels of saturated hydrocarbons (suberin and fatty acids) and phenolic compounds than in susceptible clones. These compounds could restrict the spread of O. novo-ulmi and contribute to preserving the near-normal physiological function of resistant trees when exposed to the pathogen. These results evidence common physiological responses of U. minor to flooding, drought and pathogen infection leading to xylem water disruption, leaf water stress and reduced net carbon gain. Still, seedlings of U. minor develop various mechanisms of acclimation to abiotic stresses that can play a role in surviving moderate periods of flood and drought. The chemical profile appears to be an important factor for the resistance of some genotypes of U. minor to DED. How abiotic stresses such as flooding and drought affect the capacity of resistant U. minor clones to face O. novo-ulmi is a key question that must be contemplated in future research.

Plant growth in elevated CO2 alters mitochondrial number and chloroplast fine structure

With increasing interest in the effects of elevated atmospheric CO2 on plant growth and the global carbon balance, there is a need for greater understanding of how plants respond to variations in atmospheric partial pressure of CO2. Our research shows that elevated CO2 produces significant fine structural changes in major cellular organelles that appear to be an important component of the metabolic responses of plants to this global change. Nine species (representing seven plant families) in several experimental facilities with different CO2-dosing technologies were examined. Growth in elevated CO2 increased numbers of mitochondria per unit cell area by 1.3–2.4 times the number in control plants grown in lower CO2 and produced a statistically significant increase in the amount of chloroplast stroma (nonappressed) thylakoid membranes compared with those in lower CO2 treatments. There was no observable change in size of the mitochondria. However, in contrast to the CO2 effect on mitochondrial number, elevated CO2 promoted a decrease in the rate of mass-based dark respiration. These changes may reflect a major shift in plant metabolism and energy balance that may help to explain enhanced plant productivity in response to elevated atmospheric CO2 concentrations.

Effects of elevated atmospheric CO2 concentration on leaf dark respiration of Xanthium strumarium in light and in darkness

Leaf dark respiration (R) is an important component of plant carbon balance, but the effects of rising atmospheric CO2 on leaf R during illumination are largely unknown. We studied the effects of elevated CO2 on leaf R in light (RL) and in darkness (RD) in Xanthium strumarium at different developmental stages. Leaf RL was estimated by using the Kok method, whereas leaf RD was measured as the rate of CO2 efflux at zero light. Leaf RL and RD were significantly higher at elevated than at ambient CO2 throughout the growing period. Elevated CO2 increased the ratio of leaf RL to net photosynthesis at saturated light (Amax) when plants were young and also after flowering, but the ratio of leaf RD to Amax was unaffected by CO2 levels. Leaf RN was significantly higher at the beginning but significantly lower at the end of the growing period in elevated CO2-grown plants. The ratio of leaf RL to RD was used to estimate the effect of light on leaf R during the day. We found that light inhibited leaf R at both CO2 concentrations but to a lesser degree for elevated (17–24%) than for ambient (29–35%) CO2-grown plants, presumably because elevated CO2-grown plants had a higher demand for energy and carbon skeletons than ambient CO2-grown plants in light. Our results suggest that using the CO2 efflux rate, determined by shading leaves during the day, as a measure for leaf R is likely to underestimate carbon loss from elevated CO2-grown plants.

Quantificação de biomassa e carbono da parte aérea em uma área de Mata Atlântica, na Serra da Cantareira, São Paulo

A atividade humana tem contribuído com as emissões de gases de efeito estufa (GEE) associadas, principalmente, com queima de combustíveis fósseis e mudanças no uso da terra. Assim, se faz necessário que sejam adotadas medidas visando o retardamento dos efeitos das mudanças climáticas. As florestas exercem papel essencial no balanço de carbono principalmente por funcionarem como sumidouros de CO2. Por outro lado, se desmatadas, promovem emissões e liberam parte do carbono estocado. A quantidade de biomassa florestal e o teor de carbono podem variar em função do tipo florestal, bem como de sua localização. Entretanto, fator importante diz respeito à confiabilidade dos dados mensurados neste tipo de pesquisa. A biomassa e o carbono da parte aérea podem ser determinados via método destrutivo, ou estimados via método não destrutivo. A construção do Rodoanel Mário Covas trecho norte e a supressão de uma área de Mata Atlântica possibilitou a realização de estudo de biomassa da parte aérea via método destrutivo. O objetivo deste trabalho foi estudar o tamanho e forma de parcelas, a intensidade amostral, quantificar a biomassa e o carbono na parte aérea, comparar métodos destrutivos e não destrutivos para a quantificação de biomassa e carbono na parte aérea, estudar a variação da densidade básica da madeira das espécies nas diferentes classes de DAP e grupos sucessionais e comparar as medidas de altura total e DAP obtidas a campo no inventário com as medidas coletadas após o corte. O tamanho mais conveniente de parcela foi 400 m 2, com forma retangular e dimensão de 10 x 40 m. A intensidade amostral variou entre 39 e 75 unidades amostrais. A biomassa da parte aérea obtida, via método destrutivo, foi de 188,3 Mg ha-1 e o carbono, 85,1 Mg ha-1. A biomassa estimada por equações alométricas da literatura foi subestimada, quando comparada ao valor real, obtido via método destrutivo. As menores classes de DAP apresentaram as maiores densidades básicas da madeira. A densidade básica foi 0,488 g cm-3 na média das espécies. A porcentagem de carbono contida nos troncos e galhos não diferiu entre as classes de DAP. O teor de carbono foi 45,41%, na média dos troncos e galhos. Espécies pioneiras acumularam maior quantidade de biomassa e carbono nos galhos e apresentaram maior densidade básica que as não pioneiras. A utilização dos dados coletados na fase de inventário e após o corte não afetaram os valores de biomassa estimados.

PipX, the coactivator of NtcA, is a global regulator in cyanobacteria

To modulate the expression of genes involved in nitrogen assimilation, the cyanobacterial PII-interacting protein X (PipX) interacts with the global transcriptional regulator NtcA and the signal transduction protein PII, a protein found in all three domains of life as an integrator of signals of the nitrogen and carbon balance. PipX can form alternate complexes with NtcA and PII, and these interactions are stimulated and inhibited, respectively, by 2-oxoglutarate, providing a mechanistic link between PII signaling and NtcA-regulated gene expression. Here, we demonstrate that PipX is involved in a much wider interaction network. The effect of pipX alleles on transcript levels was studied by RNA sequencing of S. elongatus strains grown in the presence of either nitrate or ammonium, followed by multivariate analyses of relevant mutant/control comparisons. As a result of this process, 222 genes were classified into six coherent groups of differentially regulated genes, two of which, containing either NtcA-activated or NtcA-repressed genes, provided further insights into the function of NtcA–PipX complexes. The remaining four groups suggest the involvement of PipX in at least three NtcA-independent regulatory pathways. Our results pave the way to uncover new regulatory interactions and mechanisms in the control of gene expression in cyanobacteria.

Synthesis dataset of physical and ecosystem properties from pan-arctic wetland sites using peat core analysis

Permafrost dynamics play an important role in high-latitude peatland carbon balance and are key to understanding the future response of soil carbon stocks. Permafrost aggradation can control the magnitude of the carbon feedback in peatlands through effects on peat properties. We compiled peatland plant macrofossil records for the northern permafrost zone (515 cores from 280 sites) and classified samples by vegetation type and environmental class (fen, bog, tundra and boreal permafrost, thawed permafrost). We examined differences in peat properties (bulk density, carbon (C), nitrogen (N) and organic matter content, C/N ratio) and C accumulation rates among vegetation types and environmental classes.

Biodegradation of naphthalene-2-sulfonic acid present in tannery wastewater by bacterial isolates Arthrobacter sp 2AC and Comamonas sp 4BC

Two bacterial strains, 2AC and 4BC, both capable of utilizing naphthalene-2-sulfonic acid (2-NSA) as a sole source of carbon, were isolated from activated sludges previously exposed to tannery wastewater. Enrichments were carried out in mineral salt medium (MSM) with 2-NSA as the sole carbon source. 16S rDNA sequencing analysis indicated that 2AC is an Arthrobacter sp. and 4BC is a Comamonas sp. Within 33 h, both isolates degraded 100% of 2-NSA in MSM and also 2-NSA in non-sterile tannery wastewater. The yield coefficient was 0.33 g biomass dry weight per gram of 2-NSA. A conceptual model, which describes the aerobic transformation of organic matter, was used for interpreting the biodegradation kinetics of 2-NSA. The half-lives for 2-NSA, at initial concentrations of 100 and 500 mg/l in MSM, ranged from 20 h (2AC) to 26 h (4BC) with lag-phases of 8 h (2AC) and 12 h (4BC). The carbon balance indicates that 75-90% of the initial TOC (total organic carbon) was mineralized, 5-20% remained as DOC (dissolved organic carbon) and 3-10% was biomass carbon. The principal metabolite of 2-NSA biodegradation (in both MSM and tannery wastewater) produced by Comamonas sp. 4BC had a MW of 174 and accounted for the residual DOC (7.0-19.0% of the initial TOC and 66% of the remaining TOC). Three to ten percent of the initial TOC (33% of the remaining TOC) was associated with biomass. The metabolite was not detected when Arthrobacter sp. 2AC was used, and a lower residual DOC and biomass carbon were recorded. This suggests that the two strains may use different catabolic pathways for 2-NSA degradation. The rapid biodegradation of 2-NSA (100 mg/l) added to non-sterile tannery wastewater (total 2-NSA, 105 mg/l) when inoculated with either Arthrobacter 2AC or Comamonas 4BC showed that both strains were able to compete with the indigenous microorganisms and degrade 2-NSA even in the presence of alternate carbon sources (DOC in tannery wastewater = 91 mg/l). The results provide information useful for the rational design of bioreactors for tannery wastewater treatment.

The response of lichen growth to additions of distilled water, rainwater and water from a rock surface

Three lichen species were wetted in the field with distilled water, rainwater or water which had run off a rock surafce, during July 1974 to February 1975. The radial growth rate of Parmelia glabratula ssp. fuliginosa was not influenced by the wetting treatments. The radial growth rate of P. conspersa with the distilled water was greater than the control, rainwater and runoff treatments. The radial growth rate of Physcia orbicularis was lower with rainwater and runoff treatmentss than the control and distilled water treatment. These results may be explained by the effect of wetting on the carbon balance of the lichens and by the influence of water chemistry.

The biology of rudraksha

Rudraksha, used throughout India and Southeast Asia in religious jewellery, is the stony endocarp of a tree distributed from northern Australia to southern Nepal. This article summarizes its biology, particularly recent research on the remarkable fruit colour. The iridescent blue colour is caused by a remarkable structure an 'iridosome'. It is secreted by the epidermal cell, and is located above the plasmalemma and beneath the adaxial wall. Cellulosic layers within the iridosome constructively interfere with blue wavelengths, causing an intense colour production at 439 nm. This colour persists in senescing fruits and may enhance their dispersal. The transparency of the cuticle at longer wavelengths allows photosynthesis to occur in the fleshy green exocarp tissue, enhancing the carbon balance of the tree. More research will certainly reveal the evolution of this remarkable phenomenon, as well as the origins of the rudraksha's cultural use.

Arctic ecosystem responses to changes in water availability and warming: Short and long-term responses

Arctic soils store close to 14% of the global soil carbon. Most of arctic carbon is stored below ground in the permafrost. With climate warming the decomposition of the soil carbon could represent a significant positive feedback to global greenhouse warming. Recent evidence has shown that the temperature of the Arctic is already increasing, and this change is associated mostly with anthropogenic activities. Warmer soils will contribute to permafrost degradation and accelerate organic matter decay and thus increase the flux of carbon dioxide and methane into the atmosphere. Temperature and water availability are also important drivers of ecosystem performance, but effects can be complex and in opposition. Temperature and moisture changes can affect ecosystem respiration (ER) and gross primary productivity (GPP) independently; an increase in the net ecosystem exchange can be a result of either a decrease in ER or an increase in GPP. Therefore, understanding the effects of changes in ecosystem water and temperature on the carbon flux components becomes key to predicting the responses of the Arctic to climate change. The overall goal of this work was to determine the response of arctic systems to simulated climate change scenarios with simultaneous changes in temperature and moisture. A temperature and hydrological manipulation in a naturally-drained lakebed was used to assess the short-term effect of changes in water and temperature on the carbon cycle. Also, as part of International Tundra Experiment Network (ITEX), I determined the long-term effect of warming on the carbon cycle in a natural hydrological gradient established in the mid 90's. I found that the carbon balance is highly sensitive to short-term changes in water table and warming. However, over longer time periods, hydrological and temperature changed soil biophysical properties, nutrient cycles, and other ecosystem structural and functional components that down regulated GPP and ER, especially in wet areas.

Peatland depths and radiocarbon dates, recent decades, North America

Peatland ecosystems store about 500-600 Pg of organic carbon, largely accumulated since the last glaciation. Whether they continue to sequester carbon or release it as greenhouse gases, perhaps in large amounts, is important in Earth's temperature dynamics. Given both ages and depths of numerous dated sample peatlands, their rate of carbon sequestration can be estimated throughout the Holocene. Here we use average values for carbon content per unit volume, the geographical extent of peatlands, and ecological models of peatland establishment and growth, to reconstruct the time-trajectory of peatland carbon sequestration in North America and project it into the future. Peatlands there contain ~163 Pg of carbon. Ignoring effects of climate change and other major anthropogenic disturbances, the rate of carbon accumulation is projected to decline slowly over millennia as reduced net carbon accumulation in existing peatlands is largely balanced by new peatland establishment. Peatlands are one of few long-term terrestrial carbon sinks, probably important for global carbon regulation in future generations. This study contributes to a better understanding of these ecosystems that will assist their inclusion in earth-system models, and therefore their management to maintain carbon storage during climate change.

CO2 concentration and stable isotope ratios of three Antarctic ice cores covering the period from 149.4 - 1.5 kyr before 1950

We present new d13C measurements of atmospheric CO2 covering the last glacial/interglacial cycle, complementing previous records covering Terminations I and II. Most prominent in the new record is a significant depletion in d13C(atm) of 0.5 permil occurring during marine isotope stage (MIS) 4, followed by an enrichment of the same magnitude at the beginning of MIS 3. Such a significant excursion in the record is otherwise only observed at glacial terminations, suggesting that similar processes were at play, such as changing sea surface temperatures, changes in marine biological export in the Southern Ocean (SO) due to variations in aeolian iron fluxes, changes in the Atlantic meridional overturning circulation, upwelling of deep water in the SO, and long-term trends in terrestrial carbon storage. Based on previous modeling studies, we propose constraints on some of these processes during specific time intervals. The decrease in d13C(atm) at the end of MIS 4 starting approximately 64 kyr B.P. was accompanied by increasing [CO2]. This period is also marked by a decrease in aeolian iron flux to the SO, followed by an increase in SO upwelling during Heinrich event 6, indicating that it is likely that a large amount of d13C-depleted carbon was transferred to the deep oceans previously, i.e., at the onset of MIS 4. Apart from the upwelling event at the end of MIS 4 (and potentially smaller events during Heinrich events in MIS 3), upwelling of deep water in the SO remained reduced until the last glacial termination, whereupon a second pulse of isotopically light carbon was released into the atmosphere.