Temperature influence on the carbon isotopic composition of Orbulina universa and Globigerina bulloides (planktonic foraminifera)

Laboratory experiments with the planktonic foraminifera Orbulina universa (symbiotic) and Globigerina bulloides (nonsymbiotic) were used to examine the effects of temperature, irradiance (symbiont photosynthesis), [CO32-], [HPO42-], and ontogeny on shell d13C values. In ambient seawater ([CO32-] = 171 mmol kg-1), the d13C of O. universa shells grown under low light (LL) levels is insensitive to temperature and records the d13C value of seawater TCO2. In contrast, the d13C of high light (HL) shells increases ~0.4‰ across 15-25°C (+0.050‰/°C). This suggests that the d13C enrichment due to symbiont photosynthetic activity is temperature-dependent. A comparison of HL O. universa grown in elevated [CO32-] seawater with ambient specimens shows that temperature does not affect the slope of the d13C/[CO32-] relationship previously described [Spero et al., 1997]. The d13C of G. bulloides shells decreases across the 15-24°C temperature range and d13C:temperature slopes decrease with increasing shell size (-0.13, -0.10, and -0.09‰/°C in 11- 12-, and 13-chambered shells, respectively). The pattern of lower d13C values at higher temperatures likely results from the incorporation of more respired CO2 into the shell at higher metabolic rates. The d13C of HL O. universa increases with increased seawater [HPO42-]

Insight into deep-sea life – Cibicidoides pachyderma substrate and pHdependent behaviour following disturbance

Most palaeo-deep-water reconstructions are based on geochemical information stored in the calcareous shells of Cibicidoides species but hardly anything is known about their life cycle, population dynamics or ecology. The number of specimens of a single Cibicidoides species can locally be very limited and species may be lacking completely during certain intervals in the geological past. As a consequence, geochemical analyses are often carried out on lumped Cibicidoides spp. assuming that they share the same epizoic to epifaunal habitat and precipitated their shell in comparable offsets to surrounding bottom water mass properties. However, there is a growing body of evidence that particularly Cibicidoides pachyderma and its morphotypes C. mundulus and C. kullenbergi, may not be reliable bottom water recorders. We have recently developed aquaria that allowed, for the first time, observations of Cibicidoides pachyderma var. C. mundulus under in situ pressure and temperature. Experiments were carried out with and without artificial sediments to simulate soft sediments and rocks, respectively. Seawater was set to pH 8 and pH 7.4 to simulate more or less particulate carbon export or more or less ventilation of bottom water. Our experiments demonstrate that C. mundulus may opt for an epifaunal or an infaunal habitat depending on elapsed time following physical disturbance, pH, current activity, the availability of sediments and growth. The specimen's initial response following transfer from atmospheric pressure into the high-pressure aquaria was to immerse into the sediment or to cover more or less parts of the test with aggregated sediments or algae. However, within 24 h a strong rheotaxis became apparent and most specimens moved to sites of increased current activity under normal pH conditions (pH 8). Only few specimens remained in algae cysts or in the sediment in the pH-8 experiment. On the contrary, all specimens under pH 7.4 agglutinated a firm sediment cyst around their test and remained infaunal throughout the experimental period of three months. Independent of pH, growth was only observed in specimens that lived within an agglutinated cyst or infaunal. A solid thick cyst covered the specimens of the pH 7.4 experiment throughout the experiment and possibly restricted water exchange between the in-cyst water and the surrounding artificial bottom water mass. We suggest that a more fragile and possibly more porous sedimentary envelope may, at least temporally, have covered the infaunal specimens under pH 8 but no evidence for this was found upon termination of the experiment

Investigating the effects of growth rate and temperature on the B/Ca ratio and δ11B during inorganic calcite formation

To deconvolve the effect of growth rate and temperature on the boron partitioning into calcite and its isotope fractionation, seeded calcite precipitation experiments were performed at a constant temperature and various growth rates and at a constant growth rate and various temperatures. We show that boron partitioning increases with increasing growth rate and decreases with increasing temperature. The B isotope fractionation between calcite and B(OH)4− increases with increasing growth rate favoring the lighter B isotope for incorporation into calcite whereas no effect of temperature was observed within the temperature range investigated (12 °C to 32 °C). At the lowest temperature and growth rate δ11B of the calcite almost equals that of B(OH)4− in solution. Applying the surface entrapment model (SEMO) of Watson and Liang (1995) to our data, we demonstrate that the observed effects of temperature and growth rate on B concentration can be explained by processes in the near surface layer of the calcite crystal

Direct effects of CO2 concentration on growth and isotopic composition of marine plankton.

The assessment of direct effects of anthropogenic CO2 increase on the marine biota has received relatively little attention compared to the intense research on CO2-related responses of the terrestrial biosphere. Yet, due to the rapid air–sea gas exchange, the observed past and predicted future rise in atmospheric CO2 causes a corresponding increase in seawater CO2 concentrations, [CO2], in upper ocean waters. Increasing [CO2] leads to considerable changes in the surface ocean carbonate system, resulting in decreases in pH and the carbonate concentration, [CO2−3]. These changes can be shown to have strong impacts on the marine biota. Here we will distinguish between CO2-related responses of the marine biota which (a) potentially affect the ocean's biological carbon pumps and (b) are relevant to the interpretation of diagnostic tools (proxies) used to assess climate change on geological times scales. With regard to the former, three direct effects of increasing [CO2] on marine plankton have been recognized: enhanced phytoplankton growth rate, changing elemental composition of primary produced organic matter, and reduced biogenic calcification. Although quantitative estimates of their impacts on the oceanic carbon cycle are not yet feasible, all three effects increase the ocean's capacity to take up and store atmospheric CO2 and hence, can serve as negative feedbacks to anthropogenic CO2 increase. With respect to proxies used in palaeo-reconstructions, CO2-sensitivity is found in carbon isotope fractionation by phytoplankton and foraminifera. While CO2- dependent isotope fractionation by phytoplankton may be of potential use in reconstructing surface ocean pCO2 at ancient times, CO2-related effects on the isotopic composition of foraminiferal shells confounds the use of the difference in isotopic signals between planktonic and benthic shells as a measure for the strength of marine primary production. The latter effect also offers an alternative explanation for the large negative swings in δ13C of foraminiferal calcite between glacial and interglacial periods. Changes in [CO2−3] affect the δ18O in foraminiferal shells. Taking this into account brings sea surface temperature estimates for the glacial tropics closer to those obtained from other geochemical proxies

Insight into deep-sea life - Cibicidoides pachyderma substrate and pH-dependent behaviour following disturbance

Most palaeo-deep-water reconstructions are based on geochemical information stored in the calcareous shells of Cibicidoides species but hardly anything is known about their life cycle, population dynamics or ecology. The number of specimens of a single Cibicidoides species can locally be very limited and species may be lacking completely during certain intervals in the geological past. As a consequence, geochemical analyses are often carried out on lumped Cibicidoides spp. assuming that they share the same epizoic to epifaunal habitat and precipitated their shell in comparable offsets to surrounding bottom water mass properties. However, there is a growing body of evidence that particularly Cibicidoides pachyderma and its morphotypes C. mundulus and C. kullenbergi, may not be reliable bottom water recorders. We have recently developed aquaria that allowed, for the first time, observations of Cibicidoides pachyderma var. C. mundulus under in situ pressure and temperature. Experiments were carried out with and without artificial sediments to simulate soft sediments and rocks, respectively. Seawater was set to pH 8 and pH 7.4 to simulate more or less particulate carbon export or more or less ventilation of bottom water. Our experiments demonstrate that C. mundulus may opt for an epifaunal or an infaunal habitat depending on elapsed time following physical disturbance, pH, current activity, the availability of sediments and growth. The specimen's initial response following transfer from atmospheric pressure into the high-pressure aquaria was to immerse into the sediment or to cover more or less parts of the test with aggregated sediments or algae. However, within 24 h a strong rheotaxis became apparent and most specimens moved to sites of increased current activity under normal pH conditions (pH 8). Only few specimens remained in algae cysts or in the sediment in the pH-8 experiment. On the contrary, all specimens under pH 7.4 agglutinated a firm sediment cyst around their test and remained infaunal throughout the experimental period of three months. Independent of pH, growth was only observed in specimens that lived within an agglutinated cyst or infaunal. A solid thick cyst covered the specimens of the pH 7.4 experiment throughout the experiment and possibly restricted water exchange between the in-cyst water and the surrounding artificial bottom water mass. We suggest that a more fragile and possibly more porous sedimentary envelope may, at least temporally, have covered the infaunal specimens under pH 8 but no evidence for this was found upon termination of the experiment

Decoupled carbonate chemistry controls on the incorporation of boron into Orbulina universa

In order to fully constrain paleo-carbonate systems, proxies for two out of seven parameters, plus temperature and salinity, are required. The boron isotopic composition (δ11B) of planktonic foraminifera shells is a powerful tool for reconstructing changes in past surface ocean pH. As B(OH)4− is substituted into the biogenic calcite lattice in place of CO32−, and both borate and carbonate ions are more abundant at higher pH, it was suggested early on that B ∕ Ca ratios in biogenic calcite may serve as a proxy for [CO32−]. Although several recent studies have shown that a direct connection of B ∕ Ca to carbonate system parameters may be masked by other environmental factors in the field, there is ample evidence for a mechanistic relationship between B ∕ Ca and carbonate system parameters. Here, we focus on investigating the primary relationship to develop a mechanistic understanding of boron uptake. Differentiating between the effects of pH and [CO32−] is problematic, as they co-vary closely in natural systems, so the major control on boron incorporation remains unclear. To deconvolve the effects of pH and [CO32−] and to investigate their impact on the B ∕ Ca ratio and δ11B, we conducted culture experiments with the planktonic foraminifer Orbulina universa in manipulated culture media: constant pH (8.05), but changing [CO32−] (238, 286 and 534 µmol kg−1 CO32−) and at constant [CO32−] (276 ± 19.5 µmol kg−1) and varying pH (7.7, 7.9 and 8.05). Measurements of the isotopic composition of boron and the B ∕ Ca ratio were performed simultaneously using a femtosecond laser ablation system coupled to a MC-ICP-MS (multiple-collector inductively coupled plasma mass spectrometer). Our results show that, as expected, δ11B is controlled by pH but it is also modulated by [CO32−]. On the other hand, the B ∕ Ca ratio is driven by [HCO3−], independently of pH. This suggests that B ∕ Ca ratios in foraminiferal calcite can possibly be used as a second, independent, proxy for complete paleo-carbonate system reconstructions. This is discussed in light of recent literature demonstrating that the primary relationship between B ∕ Ca and [HCO3−] can be obscured by other environmental parameters

Exploring foraminiferal Sr/Ca as a new carbonate system proxy

In present day paleoclimate research one of the most pressing challenges is the reconstruction of atmospheric CO2 concentrations. A variety of proxies for several components of the marine inorganic carbon system have been developed in this context (e.g. B isotopes, B/Ca, U/Ca) to allow reconstruction of past seawater pH, HCO3− and CO32− and thereby facilitate estimates of past atmospheric pCO2. Based on culture experiments using the benthic foraminifera Ammonia sp. we describe a positive correlation between Sr/Ca and the carbonate system, namely DIC/bicarbonate ion concentration. Foraminiferal Sr/Ca ratios provide potentially additional constraints on the carbonate system proxy, because the analysis of foraminiferal carbonate Sr/Ca is straightforward and not easily contaminated. Applying our calibration to a published dataset of paleo-Sr/Ca suggests the validity of Sr/Ca as a carbonate system proxy. Furthermore, we explore how our data can be used to advance conceptual understanding of the foraminiferal biomineralization mechanism

Effect of different seawater Mg2Â + concentrations on calcification in two benthic foraminifers

Magnesium, incorporated in foraminiferal calcite (Mg/CaCC), is used intensively to reconstruct past seawater temperatures but, in addition to temperature, the Mg/CaCC of foraminiferal tests also depends on the ratio of Mg and Ca in seawater (Mg/CaSW). The physiological mechanisms responsible for these proxy relationships are still unknown. This culture study investigates the impact of different seawater Mg2 + on calcification in two benthic foraminiferal species precipitating contrasting Mg/{CaCC}: Ammonia aomoriensis, producing low-Mg calcite and Amphistegina lessonii, producing intermediate-Mg calcite. Foraminiferal growth and test thickness were determined and, Mg/Ca was analyzed using Laser Ablation-Inductively Coupled Plasma-Mass Spectrometry ({LA}-{ICP}-{MS}). Results show that at present-day seawater Mg/{CaSW} of {\textasciitilde} 5, both species have highest growth rates, reflecting their adaptation to modern seawater element concentrations. Test thickness is not significantly affected by different Mg/{CaSW}. The relationship between Mg/{CaSW} and Mg/{CaCC} shows a distinct positive y-axis intercept, possibly reflecting at least two processes involved in foraminiferal biomineralization. The associated Mg partition ({DMg}) changes non-linearly with increasing Mg/{CaSW}, hence suggesting that the {DMg} is best described by an exponential function approaching an asymptote