Microsensor studies of photosynthesis and respiration in larger symbiotic foraminifera. I The physico-chemical microenvironment of Marginopora vertebralis, Amphistegina lobifera and Amphisorus hemprichii

 The physico-chemical microenvironment of larger benthic foraminifera was studied with microsensors for O₂, CO₂, pH, Ca²⁺ and scalar irradiance. Under saturating light conditions, the photosynthetic activity of the endosymbiotic algae increased the O₂ up to 183% air saturation and a pH of up to 8.6 was measured at the foraminiferal shell surface. The photosynthetic CO₂ fixation decreased the CO₂ at the shell down to 4.7 μM. In the dark, the respiration of host and symbionts decreased the O₂ level to 91% air saturation and the CO₂ concentration reached up to 12 μM. pH was lowered relative to the ambient seawater pH of 8.2. The endosymbionts responded immediately to changing light conditions, resulting in dynamic changes of O₂, CO₂ and pH at the foraminiferal shell surface during experimentally imposed light–dark cycles. The dynamic concentration changes demonstrated for the first time a fast exchange of metabolic gases through the perforate, hyaline shell of Amphistegina lobifera. A diffusive boundary layer (DBL) limited the solute exchange between the foraminifera and the surrounding water. The DBL reached a thickness of 400–700 μm in stagnant water and was reduced to 100–300 μm under flow conditions. Gross photosynthesis rates were significantly higher under flow conditions (4.7 nmol O₂ cm⁻³ s⁻¹) than in stagnant water (1.6 nmol O₂ cm ⁻³ s⁻¹), whereas net photosynthesis rates were unaffected by flow conditions. The Ca²⁺ microprofiles demonstrated a spatial variation in sites of calcium uptake over the foraminiferal shells. Ca²⁺ gradients at the shell surface showed total Ca²⁺ uptake rates of 0.6 to 4.2 nmol cm⁻² h⁻¹ in A. lobifera and 1.7 to 3.6 nmol cm⁻² h⁻¹ in Marginopora vertebralis. The scattering and reflection of the foraminiferal calcite shell increased the scalar irradiance at the surface up to 205% of the incident irradiance. Transmittance measurements across the calcite shell suggest that the symbionts are shielded from higher light levels, receiving approximately 30% of the incident light for photosynthesis. Received: 6 July 1999 / Accepted: 28 April 2000     

Symbiont photosynthesis increases both respiration and photosynthesis in the symbiotic sea anemone Anemonia viridis

Dark respiration of the symbiotic sea anemone Anemonia viridis (Forskäl) was observed to increase by 34% when anemones were exposed to hyperoxic sea water (150% oxygen saturation) overnight, and by 39% after exposure to 6 h in the light at a saturating irradiance of 300 E m^-2 s^-1 at normoxia (100% oxygen saturation). No increase due to light stimulation was observed in aposymbiotic control anemones. In darkness, the oxygen concentration of the coelenteric fluid was hypoxic. However, within 10 min of anemones being illuminated, coelenteric fluid was hyperoxic, and it remained elevated throughout a 12 h light period. When measured over a 24 h period (12 h light: 12 h dark), the dark respiration rate increased gradually over the first 6 h of the light period until it was 35% above the dark night-time resting rate. It remained elevated throughout the remaining light period and for 2 h into the following dark period, after which it fell back to the resting rate. Gross photosynthesis (P ^gross) increased significantly when anemones were exposed to either hyperoxia (150% oxygen saturation) or 300 E m^-2 s^-1 at normoxia. This increase was not observed when symbiotic anemones were illuminated at a low-light intensity of 100 E m^-2 s^-1. The results of this study suggest that respiration in the dark is limited by oxygen diffusion and that normal respiration is restored in the daytime by utilisation of the oxygen released by photosynthesis. Furthermore, it appears that the increased respiration following exposure to high-light intensities provides a CO₂-rich intracellular environment which further enhances the photosynthetic rate of the zooxanthellae.     

Competition for inorganic carbon between photosynthesis and calcification in the symbiont-bearing foraminifer Amphistegina lobifera

The feeding behaviour of the subtropical, intertidal crab Gaetice depressus (He Haan) has been investigated. This species was found to be an omnivorous scavenger capable of ingesting both animal and plant tissues which are passed to the mouth using the chelipeds. Furthermore, the mouth-parts themselves are modified and bear elongated setae, notably on the third maxillae. The net-like structures which the setae give rise to can be repeatedly flailed through the seawater in unison enabling the crabs to feed on suspended material. Beating frequency decreased with crab size but rose with increasing suspension concentration. Experiments demonstrated that suspension feeding can contribute significantly to nutrient uptake and direct observations confirmed that the crabs use this method in situ. The crabs are also able to use the setose appendages to sweep deposited material from the underlying substrate towards the mouth, presumably for nutritive purposes. The significance of these different feeding methods is discussed.     

Temperature acclimation of respiration and photosynthesis in the brown algaLaminaria saccharina

Sporophytes of the brown algaLaminaria saccharina (L.) Lamour grown at 15°C contained significantly more chlorophylla (chla) than did similar plants grown at 5°C. The increase in chla in 15°C plants was due to increased numbers of photosystem II reaction centes, and possibly to increased photosynthetic unit size, compared with 5°C plants. These changes were associated with increased values (photosynthetic efficiencies) in 15°C-grownL. saccharina relative to 5°C-grown plants. The changes in together with reduced respiration rates allowed 15°C-grownL. saccharina to achieve net photosynthesis and light-saturated photosynthesis at a lower photon fluence rate (PFR) than 5°C plants when both groups were assayed at the same temperature (15°C). The photon fluence rates necessary to reach the compensation point and achieve light-saturated photosynthesis (I _c andI _k , respectively) increased with increasing incubation temperature inL. saccharina grown at both 5 and 15°C. However, acclimation responses to growth temperature compensated for the short-term effect of temperature onI _c andI _k . Consequently, plants grown at 5 and 15°C were able to achieve similar rates of light-limited photosynthesis, and similarI _c andI _k values at their respective growth temperatures. These responses are undoubtedly important for perennial seaweeds such asL. saccharina, which frequently grow in light-limited habitats and experience pronounced seasonal changes in water temperature.Please address all correspondence and requests for reprints to I.R. Davison     

Effects of temperature on photosynthesis and respiration in hermatypic corals

Photosynthesis and respiration rates of the reef corals Pocillopora damicornis (Linn.), Montipora verrucosa (Lamarck), Porites compressa Dana and Fungia scutaria Lamarck were measured under controlled temperatures. Results indicate that coral metabolism is closely adapted to ambient temperature conditions. Tropical corals measured at Enewetak, Marshall Islands, showed greater primary production compared to maintenance requirements at elevated temperatures than did subtropical varieties of the same species in Hawaii. Photosynthesis: respiration (P:R) ratios were significantly and negatively related with temperature between 18° and 31°C for all Hawaiian corals, whereas at Enewetak this ratio generally showed a curvilinear relationship for this temperature range. Extrapolations of P:R regressions on temperatures to a value of 2.0 (estimated as a minimum required for long-term functional autotrophy) coincide for Hawaiian specimens with published upper lethal temperatures. Extrapolation of P:R regressions for Enewetak specimens at temperatures above 25°C suggests lethal temperatures for these corals to be 2 to 5 C° higher than for Hawaiian corals, in good agreement with recent experimental findings. Interspecific differences in P:R temperature regressions for Hawaiian corals correlating with upper lethal temperature tolerances are described.Contribution No. 505 of the Hawaii Institute of Marine Biology.     

Factors affecting respiration and photosynthesis by the benthic community of a subtical siliceous sediment

In a shallow, subtidal, siliceous sediment, benthic microalgal biomass (g chlorophyll a cm^-3) is influenced by light and physical sediment dynamics. The microalgal community is relatively dense, despite adverse conditions (7.0 g chlorophyll a cm^-3), and is able to respond rapidly to favorable conditions. Productivity of this community is significantly correlated (P0.05) with benthic light. In addition, productivity is influenced by temperature and bottom water NH₄ ⁺ and PO₄ ^-3 concentrations, especially as the concentrations fall to levels approaching the K _s (halfsaturation constant) of the microalgal community. Metabolic activity in this environment is dependent upon a continuous supply of organic carbon. Temperature is significantly correlated with respiration rate, but other factors (e.g. biomass and organic matter supply) are important also. Community respiration responds to overlying phytoplankton productivity in the same manner as deep-water benthic environments. Bacterial enumeration using CFU (colony-forming units) does not measure accurately the number of in situ metabolically active bacteria.This research was supported by Energy Research and Development Administration Contract AT (11-1) 3279, US AEC Contract AT (11-1) GEN 10, P.A. 20 and NOAA Sea Grant No. 04-3-158-22.     

Ultrastructural examination of chamber morphogenesis and biomineralization in the planktonic foraminiferOrbulina universa

Pre-sphere, trochospiralOrbulina universa (d'Orbigny) were hand-collected between June and September, 1982 to 1984, from surface waters of the Pacific Ocean ca. 2 km off Santa Catalina Island, California, USA. The formation of a spherical chamber by this planktonic foraminifer was studied with light and electron microscopy. Chamber morphogenesis is preceded by the secretion of a cytoplasmic layer rich in mitochondria. The layer expands away from the pre-existing foraminiferal shell surface in a spherical pattern. Full cytoplasm expansion is followed by the secretion of an organic matrix (OM) approximately 200 m above the original shell surface. Cytoplasm, mitochondria and/or vesicles may play a role in OM secretion. Calcification and chamber thickening, new spine growth and pore development are documented and continue for a period of 1–7 d. Approximately 24 h prior to gamete release, the foraminiferal spines are resorbed and a veneer of calcite is deposited on the shell surface. An extensive intracellular organelle unique to foraminifera, the fibrillar system, is examined and is hypothesized to play a role in the biomineralization process. The taxonomic and functional significance of theO. universa spherical chamber is discussed with respect to other foraminiferal species.     

The relative contribution of dinoflagellate photosynthesis and stored lipids to the survivorship of symbiotic larvae of the reef-building corals

The long-distance dispersal of larvae provides important linkages between populations of reef-building corals and is a critical part of coral biology. Some coral planulae have symbiotic dinoflagellates (Symbiodinium spp.) that probably provide energy in addition to the lipids provisioned within the egg. However, our understanding of the influence of symbionts on the energy metabolism and survivorship of planulae remains limited. This study examines the relative roles of symbiotic dinoflagellate photosynthesis and stored lipid content in the survivorship of the developing stages of the corals Pocillopora damicornis and Montipora digitata. We found that survivorship decreased under dark conditions (i.e. no photosynthetic activity) for P. damicornis and M. digitata at 31 and 22 days after release/spawning, respectively. The lipid content of P. damicornis and M. digitata planulae showed a significant decrease, at a higher rate, under dark conditions, when compared with light conditions. When converted to energy equivalents, the available energy provided by the depletion of lipids could account for 41.9 and 84.7% of larval metabolism for P. damicornis (by day 31) and 38.4 and 90.1% for M. digitata (by day 21) under light and dark conditions, respectively. This finding indicates that not all energy requirements of the larvae are met by lipids: energy is also sourced from the photosynthetic activities of the symbiotic dinoflagellates within these larvae, especially under light conditions. In addition, the amounts of three main lipid classes (wax esters, triglycerides, and phospholipids) decreased throughout the experiment in the planulae of both species, with the wax ester content decreasing more rapidly under dark conditions than under light conditions. The observations that the planulae of both species derive considerable amounts of energy from wax esters, and that symbiotic dinoflagellates enable larvae to use their stores at lower rates, suggested that symbiotic dinoflagellates have the potential to extend larval life under light conditions.     

Effects of light intensity on photosynthesis and dark respiration in six species of marine phytoplankton

Using an oxygen polarographic electrode, the shapes of photosynthetic curves and the effects of light on dark respiration in 6 species of marine phytoplankton wer examined. The species used were Skeletonema costatum, Ditylum brightwellii, Cyclotella nana (Thalassiosira pseudonana) (all Bacillariophyceae), Dunaliella tertiolecta (Chlorophyceae), Isochrysis galbana (Haptophyceae), and Gonyaulax tamarensis (Dinophyceae). A hysteresis was observed in all species examined with respect to increasing and decreasing light. Compensation light intensities varied by over 4 orders of magnitude, suggesting that the 1% light depth is an ambiguous measure of the euphotic zone. The data suggest that dark respiration accounts for ca. 25% of gross photosynthesis, but is species-dependent. In addition, respiration versus cell size does not describe an inverse exponential function over the size scales examined.This research was performed under the auspices of the United States Energy Research and Development Administration under Contract No. EY-76-C-02-0016.     

青藏高原高寒草地植物光合与土壤呼吸研究进展

青藏高原是全球变化研究的热点地区之一,而高寒草地碳过程研究一直都是全球变化研究中的重要课题.简要综述了近年来青藏高原高寒草地的重要碳过程-植物光合与土壤呼吸的研究进展和不足,并展望未来的研究热点.主要包括高寒草地植物叶片光合生理特征及其对强太阳辐射、低CO2分压、低温等高寒生境特征的适应,植物群落光合生理特征,植物生产力与植被碳库状况;高寒草地土壤呼吸日变化与季节变化特征及其影响因子分析,土壤有机碳(SOC)储量及其周转研究等.目前高寒草地的植物光合生理研究多基于叶片水平,群落和生态系统水平研究较少;已有的土壤呼吸研究多为对总土壤呼吸的研究,而将总呼吸区分为根呼吸、根际微生物呼吸、植物凋落物的微生物呼吸、有根区的土壤有机质分解以及无根区的土壤有机质分解等仍然是目前的研究难点.并且目前高寒草地植物光合与土壤呼吸研究缺乏学科交叉,多为相互独立、单一的植物生理学、生态学或土壤学研究.因此有关植物光合与土壤呼吸的整合性研究将是未来该区域碳循环研究的一个新方向.有关这一独特地域单元碳过程整合研究的大量开展,必将进一步提高人们对青藏高原区域碳循环机理的认知水平.     

Metabolic studies on thiobiotic free-living nematodes and their symbiotic microorganisms

The marine, free-living Stilbonematinae (Nematoda: Desmodoridae) are remarkable for the ectosymbiotic, prokaryotic microorganisms that populate their entire body surface. These nematodes occur in sulfidic sediments in the microoxic zone just above the sulfide maximum. Several facts point to a chemolithotrophic, sulfide oxidizing nature of the microorganisms. The oxygen uptake of three species was measured with and without their microbial coat using Cartesian and Gradient Diver microrespirometry in February 1989 at Carrie Bow Cay (Belize Barrier Reef). Symbiont-free stilbonematids exhibited constant and uniform oxygen uptake rates over several hours; rates which are significantly lower than those of oxyphilic nematodes. Freshly extracted stilbonematids, with intact bacterial coats, consumed significantly more oxygen than symbiont-free worms in the first 3 h of measurement. While the rates of aposymbiotic worms were more or less constant over time, the rates of symbiont-carrying worms exhibited a conspicuous drop during prolonged respiration. InStilbonema sp., symbiont carrying individuals kept under oxygenated conditions for more than 12 h had a respiration rate similar to those of aposymbiotic specimens. When such worms were re-incubated in sulfide-enriched seawater the respiration rate was significantly elevated. The possibility of recharging the oxygenated symbiosis system via sulfide-uptake is seen as an indication that storage of reduced sulfur compounds, or reserve substances synthetized in the presence of sulfide, play a decisive role in the metabolisms of the symbiotic bacteria. Migration of nematodes between sulfidic and oxidized sediment-layers are, most likely, the key to understanding the success of this nematode-bacteria symbiosis.Please address all correspondence and requests for reprints to Professor J. Ott     

Plasmatic resistance and rate of respiration and photosynthesis of Zostera marina at different salinities and temperatures

Zostera marina L. was studied at the Izembek Lagoon, Alaska Peninsula. Two morphologically different forms, tidepool and subtidal, can be distinguished. Both show a high tolerance to different salinities and temperatures. The plasmatic resistance was found in a range of distilled H₂O up to 3.0 seawater (24 h) and between-6° and 34°C (12 h). Within these resistance limits, the photosynthesis, which has its maximum in normal (1.0) seawater, decreases nearly to zero not only in distilled H₂O but even in 2.0 seawater, and increases with the temperature in the tidepool form up to 35°C, but in the subtidal form up to 30°C only. At higher temperatures photosynthesis declines sharply in both forms. Respiration has its minimum in distilled H₂O and at 0°C and increases with increasing salinity and temperature.Dedicated to Prof. Dr. Dr. h. c. mult. K. Mothes, Halle, Germany (DDR) on the occasion of his 70th birthday.     

Daily photosynthesis,respiration, and carbon budgets in a tropical marine jellyfish (Mastigias sp.)

From measured diel photosynthesis and respiration rates, using oxygen electrodes, estimates of carbon flux between symbiotic algae (zooxanthellae) and host animal are presented for the marine scyphomedusan Mastigias sp. from a marine lake in Palau, Western Caroline Islands, during February and March 1982. The carbon budgets calculated for these lake medusae indicate that carbon fixed photosynthetically by zooxanthellae and made available to the host may satisfy up to 100% of the host's daily metabolic carbon demand (CZAR). The stable carbon isotope (¹³C) signature of the mesogleal carbon of lake Mastigias sp. was close to that of the zooxanthellae, supporting the interpretation that while these medusae may feed holozoically, some of their carbon comes from their symbionts. The diel photosynthesis, respiration, and preliminary estimates of carbon budgets of three individuals of another ecotype of Mastigias sp. collected from nearby oceanic lagoons are also given. Photosynthesis of lagoon medusae was generally greater than that for lake medusae of similar size, and lagoon medusae were phototrophic with respect to carbon, with commensurately greater CZAR values. Carbon translocated from the symbiotic algae also may contribute to the growth requirements of both lake and lagoon medusae. From carbon flux data, the lake jellyfish were estimated to contribute about 16% to the total primary productivity of their marine lake habitat.     

森林生态系统中土壤呼吸研究进展

土壤呼吸是土壤微生物活性和土壤肥力的一个重要指标，是土壤碳流通的一个重要过程，也是陆地生态系统碳循环的一个关键部分，对研究全球变化有非常重要影响。文章综述了森林生态系统土壤呼吸的各种测量方法，比较了静态气室法和动态气室法的优缺点，认为动态红外气体分析法是最可靠的方法之一；探讨了影响土壤呼吸速率的各种因素，指出在各生物和非生物因素中，温度对土壤呼吸的影响最大；最后提出了土壤呼吸研究过程中存在的一些问题及今后的发展方向。     

增强的UV—B辐射对黄瓜（Cucumis sativus）不同叶位叶片生长、光合作用和呼吸作用的影响

在植物生长室中,黄瓜植株第1片真叶出现后,用人工UV－B光源照射60d,测定植物各叶位叶片的生长和生理活动。结果表明,UV－B辐射条件下,植物出叶时间被延迟,叶面积和叶干重下降,降幅与叶位高低正相关;叶片含水量降低,老龄叶片（1叶,下位叶）和幼龄叶片（第5叶,上位叶）的水分降幅均高于成年叶片（第3叶,中位叶）;叶片的伸展速度,叶片数目以及单叶面积减少,致使黄瓜总叶面积下降;植株节间长度缩短,是植株矮化的重要原因;根、茎、叶等器官之间的相关生长变化不大,叶片生长在其中起重要的协调作用。UV－B降低Pn和EAQE,对光合的抑制程度随叶位升高而增加,UV－B辐射后,黄瓜叶片的光呼吸显著提高,增幅与叶片发育阶段有关,UV－B对黄瓜第1叶的暗呼吸没有影响,第2、3叶略微下降,第4叶显著升高,分析认为,植株矮化叶面积减少有利于植物适应UV－B辐射;水分含量和光合作用减少、呼吸作用增强是黄瓜生长受抑制的生理基础。图2表2参18     

Experimental ecology of the temperate scleractinian coral Astrangia danae I. Partition of respiration,photosynthesis and calcification between host and symbionts

Calcification, photosynthesis and respiration of the scleractinian coral Astrangia danae were calculated from the changes in total alkalinity, pH, calculated total CO₂, and oxygen concentration produced by colonies incubated in glass jars. A correction for changes in ammonia, nitrate and nitrite was taken into account and the method evaluated. The fluxes of oxygen and CO₂ were highly correlated (r=0.99). The statistical error of alkalinity determinations was less than 10% of the changes observed in the slowest calcifying samples. Metabolism of polyparium alone was estimated by difference after removal of tissue and reincubation of bare corallum. Zooxanthellae concentration in the polyps was obtained from cell counts made on homogenates of polyp tissue. The calculated photosynthetic rate of the zooxanthellae in vivo was 25 mol O₂ (10⁸ cell)^-1 h^-1 at a light intensity of 120 Ein m^-2 s^-1. In corals having 0.5x10⁹ zooxanthellae/dm² of colony area up to 8% of the total photosynthesis was attributed to the corallum microcosm. Polyp respiration, photosynthesis, and CaCO₃ uptake rates were all much higher than rates previously reported from A. danae, apparently because in these experiments the organisms were better fed. This increased photosynthesis in turn enhanced calcification still further. The symbiosis therefore appears to provide a growth advantage even to fed corals, under the conditions of these experiments.     

The size and function of the internal inorganic carbon pool of the foraminifer Amphistegina lobifera

The existence of an internal inorganic carbon pool in the perforate foraminifer Amphistegina lobifera, as suggested recently (ter Kuile and Erez 1987), has been established by direct measurements using a new ¹⁴C tracer method. The imperforate species Amphisorus hemprichii does not contain such a pool. The size of the pool in A. lobifera is proportional to its calcification rate and approximately equals the amount of carbon incorporated into the skeleton during 24 h. Time course experiments show that inorganic carbon (Ci) is photoassimilated at constant rates by the algal symbionts, that the pool is filled to maximum capacity in ca. 24 h, and that Ci incorporation into the skeleton starts only after the pool is filled up. During the chase phase of pulsechase experiments, all ¹⁴C initially residing in the pool is transferred to the skeleton, indicating that the pool serves for calcification and not for photosynthesis. Uptake of Ci into the pool occurs only in the light, indicating that energy may be required for this process. Furthermore, calculations of the Ci concentration inside the pool suggest that it is higher by 2 to 3 orders of magnitude compared to seawater concentration, suggesting that its accumulation is an energy dependent process.     

Mesocosm studies of DCMU-enhanced fluorescence as a measure of phytoplankton photosynthesis

When measurements of in-vivo fluorescence are used to estimate photosynthesis in the field, the marked temporal and spatial variations in phytoplankton populations, and their nutrient and light histories, have produced varied results. Natural phytoplankton populations in large, flow-through mesocosms with different controlled nutrient and sewage sludge additions were sampled weekly from June to September 1984. Good correlations were observed between the increase in fluorescence upon the addition of DCMU (F) and both in-situ production and the parameters (P_m and ) of the photosynthesis-irradiance curve for these phytoplankton populations. Good correlations were also obtained between DCMU-enhanced fluorescence (F_DCMU) and chlorophyll a concentrations. The relationship between F and in-situ ¹⁴C production was consistent among mesocosms even in the face of major shifts from diatom-dominated to dinoflagellate-dominated populations. On the other hand, the F_DCMU:Chl a relationship was significantly different between mesocosms and related to species composition. It was concluded that F offers the possibility of rapidly and accurately indexing both in-situ production and the photosynthetic capacity of mixed phytoplankton populations.This study was supported by EPA cooperative agreement 810265-03 and NOAA grant NA-83-ABD-00008     

Effects of symbiont elimination and reinfection on the life processes of the planktonic foraminifer Globigerinoides sacculifer

Laboratory experiments were carried out to determine the influence of symbiotic dinoflagellates (zooxanthellae) on the shell growth, longevity, and reproductive potential of Globigerinoides sacculifer (Brady). Its symbionts were eliminated by 72-h treatment with a photosynthetic inhibitor (DCMU). Symbiont elimination resulted in earlier gametogenesis (shortened survival time) and smaller shell sizes of G. sacculifer when compared to untreated foraminifera grown in sea water. Individuals kept in continuous darkness in untreated sea water also exhibited early gametogenesis, short survival times and small shell sizes. Aposymbiotic foraminifera formed on the average one or two chambers fewer per individual and their rate of shell size increase is slower than symbiont-bearing foraminifera. Symbionts were lysed within perialgal vacuoles of G. sacculifer when subjected to DCMU treatment or kept in continuous darkness. One DCMU-treated group was reinfected with symbionts from crushed G. sacculifer donors. Soon after reinfection, these foraminifera resumed a shell growth rate and exhibited developmental stages that were nearly equivalent to those of untreated individuals, as deduced from their shell size, frequency of sac-like chambers, rate of gametogenesis, and survival time. Our experiments indicate that the symbionts aid in calcification and that elimination of symbionts triggers gametogenesis, thus shortening the life span of the foraminiferal host. The results imply that shell growth in symbiont-bearing planktonic foraminifera occurs mainly in the euphotic zone and that they do not survive for long periods below it.