Factors affecting the respiration of rabbit sperm measured with an oxygen electrode

The respiration of rabbit sperm was measured by a Clarke type electrode which has two advantages over the conventional Warburg technique, greater sensitivity, and no necessity for a carbon dioxide-free atmosphere. It was not necessary to resaturate the sample chamber of the oxygen monitor with air, down to about 30% desaturation.Rabbit seminal plasma had a measurable oxygen uptake (0.6 μl/hr/ml) but this was much less than for human seminal plasma (4.3 μl/hr/ml). Hoderate dilution of the sperm and storing the semen at 0°C after slow cooling had no effect on oxygen uptake. Unlike those of most other species, rabbit sperm were also little affected by deliberate exposure to cold shock and the respiration before and after rapid cooling to 0°C was about the same. On the other hand very brief exposure of rabbit sperm to 65°C abolished motility and greatly reduced the respiration rate. Bicarbonate (6 mM) stimulated the oxygen uptake of freshly collected samples of rabbit sperm after washing. Increasing the phosphate concentration of the medium to 80 mM did not greatly depress the oxygen uptake.     

Are altitudinal limits of equatorial stream insects reflected in their respiratory performance?

1. We measured respiration of the larvae of aquatic insects from streams in the Ecuadorian Andes in relation to oxygen saturation at 5, 8, 11, 14 and 17 °C. Polycentropus (Polycentropodidae), Lachlania (Oligoneuriidae), Anchytarsus (Ptilodactylidae) and Anacroneuria (Perlidae) represented genera absent from the highest altitudes, reaching 2720, 2930, 3120, 3450 m a.s.l., respectively, while Claudioperla (Gripopterygidae) and Anomalocosmoecus (Limnephilidae) occurred only above 2900 m a.s.l. Our purpose was to determine whether natural altitudinal limits were reflected in physiological critical points on respiration versus oxygen curves and by the effect of temperature on the ability to oxy‐regulate. 2. For all six genera, respiration was affected by oxygen saturation and temperature. Respiration (mg O₂ g⁻¹ AFDM h⁻¹) at 70% oxygen saturation (Michaelis–Menten fitted) varied from 2.6 to 7.6 between genera at 17 °C, and from 1.3 to 2.5 at 5 °C. Q₁₀ values for this temperature interval ranged 1.5–2.9 (mean 2.3). The two “high‐altitude” genera had higher respiration rates at low temperature and oxygen saturation, and their respiration rate saturated at lower temperatures, than three of the four “low‐altitude” genera. 3. The oxy‐regulatory capacity (critical points and initial decrease in respiration versus oxygen regressions) varied among genera and was affected by temperature. Lachlania, Claudioperla and Anomalocosmoecus had a higher ability to oxy‐regulate at low than at high temperatures, Anacroneuria was not clearly affected by temperature, while Polycentropus and Anchytarsus had a greater oxy‐regulatory capacity at high than at low temperature. These results indicate that the ability to oxy‐regulate is related to the temperature (altitude) at which species naturally occur. 4. Upper altitudinal limits of the six genera were not reflected in their respiratory performance, because all genera had critical minima of temperature and oxygen saturation much lower than those occurring at the limits of their natural distribution. So, the altitudinal limit could not be attributed to absolute short‐term physiological tolerance of low temperature and oxygen concentration. 5. Multiple regressions (based on respiration experiments and previously obtained relationships between water temperature, oxygen saturation and altitude) were used to predict how respiration rates should vary with altitude. At the upper limit of the four “low‐altitude” genera, respiration rates were 50–68% of those predicted at the centre of the range. With an arbitrary increase of 400 m above the actual limit, the effect of temperature would be a 13% decrease, and that of oxygen a 2% decrease, in respiration rate of Polycentropus, Lachlania and Anacroneuria, while respiration in Anchytarsus would be reduced by 5% by both factors. 6. It seems that, while the immediate decrease in respiration with increased altitude is caused mainly by a decrease in temperature, the long‐term survival of a species at given altitudes might be more affected by oxygen saturation. Further quantitative and long‐term studies on survival and recruitment in populations and communities are needed to determine the importance of temperature and oxygen for altitudinal limits of aquatic insects.     

Simulation of oxygen transfer in microbial slimes

A model based on diffusion mechanisms accounts for oxygen transfer from flowing nutrient medium into a film of organisms in slime. Profiles of dissolved oxygen concentration are generated at distances downstream from the start of the slime. In these studies, the nutrient medium was assumed rich, thus the rate of oxygen transport limited microbial respiration. Parameter sensitivity tests were performed for flow rate, oxygen concentration in the medium, and mass transfer coefficients.     

Germination and Dormancy of Reed Canary-Grass Seeds (Phalaris arundinacea)

Effects of various chemical and physical factors on the germination of several seed lots of reed canary-grass (Phalaris arundinacea L.) have been studied. Germination at the optimum constant temperatures of 24 to 27°C was significantly stimulated by the following treatments: moist chilling in light, red light given during the first 3 days of imbibition, three 2-h periods at 12°C given during the second day of imbibition, ethylene, increased oxygen tension and soaking in aerated water for 4 days. Dry storage at 20–30°C had no effect on the germination ability of the seeds. No significant quantities of germination inhibitors were found either in water or methanol extracts of seed dispersal units. By comparing three cultivars with various degrees of seed dormancy, respiration measurements showed that there was a significant positive correlation between oxygen uptake prior to visible germination and germination capacity. Similarly, germination-stimulating treatment significantly enhanced oxygen uptake prior to visible germination.     

Short‐term high temperature treatment reduces viability and inhibits respiration and DNA repair enzymes in Araucaria angustifolia cells

We evaluated the effect of global warming on Araucaria angustifolia (Bert.) O. Kuntze, a critically endangered native tree of Southern Brazil, by studying the effects of short‐term high temperature treatment on cell viability, respiration and DNA repair of embryogenic cells. Compared with control cells grown at 25°C, cell viability was reduced by 40% after incubation at 30 and 37°C for 24 and 6 h, respectively, while 2 h at 40 and 42°C killed 95% of the cells. Cell respiration was unaffected at 30–37°C, but dramatically reduced after 2 h at 42°C. The in vitro activity of enzymes of the base excision repair (BER) pathway was determined. Apurinic/apyrimidine endonuclease, measured in extracts from cells incubated for 2 h at 42°C, was completely inactivated while lower temperatures had no effect. The activities of three enzymes of the mitochondrial BER pathway were measured after 30‐min preincubation of isolated mitochondria at 25–40°C and one of them, uracil glycosylase, was completely inhibited at 40°C. We conclude that cell viability, respiration and DNA repair have different temperature sensitivities between 25 and 37°C, and that they are all very sensitive to 40 or 42°C. Thus, A. angustifolia will likely be vulnerable to the short‐term high temperature events associated with global warming.     

The relationship between respiration and temperature in leaves of the arctic plant Saxifraga cernua

Abstract Saxifraga cernua, a perennial herb distributed throughout the arctic and subarctic regions, shows high levels of dark respiration. The amount of respiration exhibited by leaves and whole plants at any temperature is influenced by the pretreatment temperature. Plants grown at 10°C typically show higher dark respiration rates than plants grown at 20°C. The levels of alternative-pathway respiration (or cyanide-insensitive respiration) in leaves of S. cernua grown at high and low temperatures were assessed by treating leaf discs with 0.25 mol m^?3 salicylhydroxamic acid during measurements of oxygen consumption. Alternative pathway respiration accounted for up to 75% of the total respiration. Tissues from 20°C-grown plants yielded a Q₁₀ of 3.37 for normal respiration, and of 0.97 for alternative-pathway respiration. Tissues from 10°C-grown plants yielded a Q₁₀ of 2.55 for normal respiration, and of 0.79 for alternative-pathway respiration. The alternative pathway does not appear to be as temperature sensitive as the normal cytochrome pathway. A simple energy model was used to predict the temperature gain expected from these high rates of alternative-pathway respiration. The model shows that less than 0.02°C can be gained by leaves experiencing these high respiration rates.     

Blooms of Surf-zone Diatoms Along the Coast of the Olympic Peninsula,Washington. VIII. Effect of Temperature and Oxygen Concentration on Respiration of Chaetoceros armatum in the Natural Environment

Dark respiration rates of the surf diatom species, Chaetoceros armatum, were measured in field experiments throughout a year-round study. Measurements of dissolved oxygen were made by employing the Winkler oxygen technique. Oxygen concentration in the ambient water had a significant positive effect upon respiration rate at all temperatures encountered during the period of study (4.6d̀ up to 13.8 d̀C). Above 8d̀, respiration rate increased with increasing temperature up to about 12.5d̀. Above 12.5d̀, temperature had a detrimental effect. The relationship between the concentration of dissolved oxygen and respiration rate was confirmed experimentally by artificially lowering or raising the oxygen concentration.     

Physiology of diapause in the adult bark beetle, Ips acuminatus Gyll., studied in relation to cold hardiness

The ovarioles of the bark beetle, Ips acuminatus are telotrophic. Ovarian development is suppressed at an immature stage with primary germ cells present in the germaria. Lower oxygen consumption is found in beetles during autumn and early winter, and a substantial rise in mean respiration rates occurs in the beginning of January paralleled by a resumption of pre-vitellogenesis in all females maintained either at 3°C or out of doors after return to 21°C for 2 weeks. It is concluded that I. acuminatus enters faculative diapause soon after enclosure to the adult, and that diapause is terminated by mid-winter in beetles kept for 18 weeks at either 3°C or out of doors. The specimens remain thereafter in reproductive quiescence until ovarian development can proceed.Photic cues are neither involved in the elevation of mean respiration rates, nor needed to abolish the inhibition of ovarian maturation in beetles kept at 3°C or in those returned to 21°C. However, follicle formation in ovarioles is only seen in positive phototactic females reared during “long-day” conditions, suggesting a photoperiodic regulation of the later stages of vitellogenesis.Detectable amounts of ethylene glycol are found at the beginning of November in freezing-susceptible I. acuminatus hibernating in its galleries underneath bark of Scots pine (Pinus silvestris) at 3°C. The gradual catabolism of the cryoprotective solute at 3°C through December occurred at a time when individuals achieved the competence to resume ovarian maturation during 2 weeks at 21°C, but prior to the substantial rise in their mean respiration rates. However, resumption of ovarian development in spring had no effect on the capability of outdoor beetles to enhance their supercooling capacity when subjected to sub-zero temperatures. Since the ability to respond to temperature changes occurred in post-diapause I. acuminatus as well, the maintenance of prolonged cold hardiness in specimens could not be related to diapause itself. Apparently, the ability of beetles to resynthesize ethylene glycol when a detectable level is present in the organism remains unaltered during overwintering.     

Effects of salinity and cyclic temperature on survival of two sympatric species of grass shrimp (Palaemonetes), and their relationship to natural distributions

Survival and respiration of the grass shrimp Palaemonetes pugio Holthuis and P. vulgaris (Say) from the Newport River estuary were measured after exposure to cyclic and constant winter temperatures, to rapid decreases in temperature, and to various temperature-salinity combinations. Both species were subjected to nine temperature-salinity combinations including temperature regimes of cyclic 7–13°C, constant 7° and 10°C, and salinities of 5, 20, and 35%.. Based on the laboratory and field results, the differences in physiological tolerance to winter temperatures and salinities were examined in relation to habitat partitioning by these sympatric species.Survival after continuous exposure to cyclic temperature regimes at medium to high salinities was similar to that observed for comparable constant temperatures; however, at low salinities mortality was significantly lower under the cyclic regime than under either constant regime. This suggests that cyclic temperatures may be detrimental in combination with some other stress. A rapid, transient decrease in temperature from either 7° or 10°C to 2°C had no measurable effect on survival or rate of oxygen consumption at any temperature-salinity acclimation. Neither salinity (except in areas intermittently subjected to salinities below 3 %.) nor winter temperatures appear to affect habitat partitioning in grass shrimp.     

Effect of cold shock and cooling rate on calcium uptake of ram spermatozoa

Rapid cooling (cold shock) of washed ejaculated ram sperm irreversibly reduced motility and respiration and greatly increased uptake of ⁴⁵Ca²⁺. The effect was greater as the temperature of cooling was reduced from 15°C to 0°C, and a substantial increase in sperm calcium levels was even observed after slow cooling to temperatures below 10°C. The rise in calcium uptake on freezing sperm to −79°C was not as great as that on cold shocking sperm to 0°C.Inactivation of sperm by mild heat (50°C) had no significant effect on calcium uptake but subsequent cold shock increased the sperm calcium. Reverse immobilization of sperm by low concentrations of formaldehyde significantly reduced calcium uptake on cold shock. Addition of detergents to sperm immediately reduced motility, respiration and calcium uptake of control and cold-shocked sperm to zero.     

Metabolite profiling reveals tissue- and temperature-specific metabolomic responses in thermoregulatory male florets of Dracunculus vulgaris (Araceae)

The male part of the spadix of Dracunculus vulgaris exhibits a degree of temperature regulation by inversely controlled heat production over a 20–35 °C range of tissue temperature. To clarify the effects of temperature on cellular metabolism, comparative analysis was performed using 51 metabolites from two distinct tissues (florets and pith) of thermogenic male spadices that had been temperature clamped at either 20 (to produce high respiration) or 35 °C (to produce low respiration). Principal component analysis and hierarchical clustering analysis showed that changes in metabolites in the florets, but not in the pith, were associated with temperature change. The energy charge in the florets treated at 20 °C was significantly higher than that of the florets treated at 35 °C. This indicated the presence of an increased energy-producing pathway that ultimately led to an increased level of thermogenesis at 20 °C. Intriguingly, succinate, a direct substrate for complex II in the mitochondrial respiratory chain, was the metabolite most significantly affected in our analysis, with its concentration in the florets 3.5 times higher at 20 than at 35 °C. However, the mitochondria fed with succinate showed that state 2 and 3 respirations and the capacity of the alternative and cytochrome pathways were all significantly higher at 35 than at 20 °C. Taken together, the results show that the male florets are the primary sites for temperature-induced changes in metabolomic pathways, although succinate-stimulated mitochondrial respiration, per sé, is not the control mechanism for thermoregulation in D. vulgaris.     

Growth temperature influences the underlying components of relative growth rate: an investigation using inherently fast- and slow-growing plant species

We examined the effect of growth temperature on the underlying components of growth in a range of inherently fast‐ and slow‐growing plant species. Plants were grown hydroponically at constant 18, 23 and 28 °C. Growth analysis was conducted on 16 contrasting plant species, with whole plant gas exchange being performed on six of the 16 species. Inter‐specific variations in specific leaf area (SLA) were important in determining variations in relative growth rate (RGR) amongst the species at 23 and 28 °C but were not related to variations in RGR at 18 °C. When grown at 18 °C, net assimilation rate (NAR) became more important than SLA for explaining variations in RGR. Variations in whole shoot photosynthesis and carbon concentration could not explain the importance of NAR in determining RGR at the lower temperatures. Rather, variations in the degree to which whole plant respiration per unit leaf area acclimated to the different growth temperatures were responsible. Plants grown at 28 °C used a greater proportion of their daily fixed carbon in respiration than did the 18 and 23 °C‐grown plants. It is concluded that the relative importance of the underlying components of growth are influenced by growth temperature, and the degree of acclimation of respiration is of central importance to the greater role played by NAR in determining variations in RGR at declining growth temperatures.     

The temperature responses of soil respiration in deserts: a seven desert synthesis

The temperature response of soil respiration in deserts is not well quantified. We evaluated the response of respiration to temperatures spanning 67°C from seven deserts across North America and Greenland. Deserts have similar respiration rates in dry soil at 20°C, and as expected, respiration rates are greater under wet conditions, rivaling rates observed for more mesic systems. However, deserts differ in their respiration rates under wet soil at 20°C and in the strength of the effect of current and antecedent soil moisture on the sensitivity and magnitude of respiration. Respiration increases with temperature below 30°C but declines for temperatures exceeding 35°C. Hot deserts have lower temperature sensitivity than cold deserts, and insensitive or negative temperature sensitivities were predicted under certain moisture conditions that differed among deserts. These results have implications for large-scale modeling efforts because we highlight the unique behavior of desert soil respiration relative to other systems. These behaviors include variable temperature responses and the importance of antecedent moisture conditions for soil respiration.     

The function of slime from Physarum flavicomum in the control of cell division.

A haploid cell of the myxomycete Physarum flavicomum undergoes cytokinesis, producing a large population of cells. However, after syngamy, cytokinesis no longer occurs but karyokinesis does and subsequent growth results in the formation of a diploid syncytial plasmodium. Slime, which is produced by the plasmodium but not the haploid cells, was aseptically isolated and purified, and tested for its effect as a cytokinetic regulator. Slime (a viscous, high molecular weight, acidic glycoprotein) affected cytokinesis of the haploid myxamoebae growing in pure culture in soluble media, and the effect was concentration dependent. In simple media, a slime concentration of about 6 10(-5) mug protein per cell suppressed cytokinesis about 50%, unequally inhibited the synthesis of protein, RNA, and DNA, but stimulated respiration. The biological activity of slime was not species specific and it also affected the bacterium Bacillus subtilis by inhibiting cytokinesis, stimulating oxygen uptake, and producing an aberrant cell morphology. Slime was inactivated by heat, fragmentation, and incubation with dithiothreitol, mercaptoethanol, and the proteolytic enzyme papain (EC 3.4.22.2). The inhibitory effect of slime on cell division of haploid cells could not be achieved using mucin or various polyanions. The possible role of slime in the production of the diploid syncytium is discussed.     

On the Metabolism of Tortula ruralis Following Desiccation and Freezing: Respiration and Carbohydrate Oxidation

Recovery from desiccation by Tortula ruralis (Hedw.) Gaertn., Meyer and Scherb was accompanied by an immediate, rapid increase in respiration (measured as oxygen uptake) at 25.5°C or 3.5°C. The respiratory burst was greater on rehydration of moss which had been rapidly desiccated over silica gel than that which had been more slowly desiccated in atmospheres of high relative humidity. No respiration was observed in dry moss. Dried moss which had been placed in liquid nitrogen resumed respiration on rewarming and rehydration but moss which had been frozen in the hydrated state respired to a lesser extent and showed signs of freeze damage. In the initial stages of slow drying a slight increase in respiration was noted, followed by a gradual decrease as drought became more severe. In contrast to observations made on many higher plants under drought stress, this moss did not exhibit any changes in its starch and sugar content during or following desiccation, nor were there any changes in free proline levels. Using (1-¹⁴C)-glucose and (6-¹⁴C)-glucose, the relative activities of the Embden–Meyerhof–Parnas and pentose phosphate pathways in hydrated and rehydrated moss were determined, as were the activities of specific enzymes involved in these pathways. An increased activity of the Embden–Meyerhof–Parnas pathway of glucose oxidation on rehydration of Tortula was observed. The possible significance of this latter observation is outlined.     

The effect of substrate, ADP and uncoupler on the respiration of tomato pollen during incubation in vitro at moderately high temperature

Pollen of tomato cv. Supermarmande was collected from greenhouse-grown plants at various intervals throughout the year and arbitrarily classified as of high, medium or low respiratory activity on the basis of CO₂ production during 8 h incubation in vitro at 30°C, a temperature that is considered to be moderately high for tomato fruit set. After an initial burst of respiration during the first stage of hydration at 30°C (>1 h), the respiration rate of pollen of all three categories declined, the decrease being greater in the lots with a low or medium respiratory activity than in the high category. During hydration (10 min after the start of incubation), the addition of succinate or reduced β-nicotinamide adenine dinucleotide (NADH) to the substrate increased the respiratory rate of slowly-respiring pollen more than that of fast-respiring pollen, but carbonyl cyanide 3-chlorophenylhydrazone (CCCP) and adenosine 5′-diphosphate (ADP) had less effect. After 1–4 h incubation, the respiration rate of the slow- or medium-respiring pollen lots had decreased, but was stimulated by succinate or NADH, and to a lesser degree by ADP. By 7 h, the respiration rate of all pollen lots had declined and was stimulated less by substrate, ADP or CCCP. The oxidation of NADH by tomato pollen contrasts with the failure of other pollen species to utilize this substrate; moreover, a synergistic effect of NADH and succinate was consistently observed. We conclude that the decline in respiration during incubation for up to 4 h at 30°C may reflect a lack of respiratory substrate. After 7 h, however, the decreased response to substrate indicates a loss of mitochondrial integrity or an accumulation of metabolic inhibitors. It is concluded that at 30°C (a moderately high temperature for tomato pollen), the initially high rate of respiration leads to exhaustion of the endogenous respiratory substrates (particularly in pollen with low to medium respiratory activity), but subsequently to ageing and a loss of mitochondrial activity.     

Temperature dependence and ontogenetic changes of metabolic rate of an endemic earthworm <Emphasis Type="Italic">Dendrobaena mrazeki</Emphasis>

Ontogenetic changes and temperature dependency of respiration rate were studied in Dendrobaena mrazeki, an earthworm species inhabiting relatively warm and dry habitats in Central Europe. D. mrazeki showed respiration rate lower than in other earthworm species, < 70 μl O₂ g⁻¹ h⁻¹, within the temperature range of 5–35°C. The difference of respiration rate between juveniles and adults was insignificant at 20°C. The response of oxygen consumption to sudden temperature changes was compared with the temperature dependence of respiratory activity in animals pre-acclimated to temperature of measurement. No significant impact of acclimation on the temperature response of oxygen consumption was found. The body mass-adjusted respiration rate increased slowly with increasing temperature from 5 to 25°C (Q₁₀ from 1.2 to 1.7) independently on acclimation history of earthworms. Oxygen consumption decreased above 25°C up to upper lethal limit (about 35°C). Temperature dependence of metabolic rate is smaller than in other earthworm species. The relationships between low metabolic sensitivity to temperature, slow locomotion and reactivity to touching as observed in this species are discussed.     

Effects of hypothermia on canine kidney mitochondria

The effect of hypothermia on the function of isolated dog kidney cortex mitochondria was determined with an FAD- and NAD⁺-linked substrate. In dog kidney mitochondria, temperatures of 10 °C or less suppress ADP stimulation of respiration but have little or no effect upon uncoupler, Ca²⁺ or valinomycin-K⁺ stimulation of respiration. This suggests that the adenine nucleotide translocase which catalyses the transport of ADP into the mitochondria limits the rate of respiration and generation of ATP at 10 °C in kidneys undergoing preservation. The coupling of oxidation to phosphylation, as determined by measuring the amount of ATP formed at low temperatures, indicates, however, that mitochondria are fully coupled at both 10 and 5 °C. The respiratory control index at 15 °C is greater (with pyruvate plus malate) than at 30 or 10 °C and suggests that 15 °C may be the optimum perfusion temperature for maintaining adenine nucleotide levels in the perfused kidney.     

The influence of increasing growth temperature and CO2 concentration on the ratio of respiration to photosynthesis in soybean seedlings

Using controlled environmental growth chambers, whole plants of soybean, cv. ‘Clark’, were examined during early development (7–20 days after sowing) at both ambient (≈ 350 μL L^–1) and elevated (≈ 700 μL L^–1) carbon dioxide and a range of air temperatures (20, 25, 30, and 35 °C) to determine if future climatic change (temperature or CO₂ concentration) could alter the ratio of carbon lost by dark respiration to that gained via photosynthesis. Although whole-plant respiration increased with short-term increases in the measurement temperature, respiration acclimated to increasing growth temperature. Respiration, on a dry weight basis, was either unchanged or lower for the elevated CO₂ grown plants, relative to ambient CO₂ concentration, over the range of growth temperatures. Levels of both starch and sucrose increased with elevated CO₂ concentration, but no interaction between CO₂ and growth temperature was observed. Relative growth rate increased with elevated CO₂ concentration up to a growth temperature of 35 °C. The ratio of respiration to photosynthesis rate over a 24-h period during early development was not altered over the growth temperatures (20–35 °C) and was consistently less at the elevated relative to the ambient CO₂ concentration. The current experiment does not support the proposition that global increases in carbon dioxide and temperature will increase the ratio of respiration to photosynthesis; rather, the data suggest that some plant species may continue to act as a sink for carbon even if carbon dioxide and temperature increase simultaneously.     

Light respiration by subtropical seaweeds

Here, we report the first‐ever measurements of light CO₂ respiration rate (CRR) by seaweeds. We measured the influence of temperature (15–25°C) and light (irradiance from 60 to 670 μmol · m^?2 · s^?1) on the light CCR of two subtropical seaweed species, and measured the CRR of seven different seaweed species under the same light (150 μmol · m^?2 · s^?1) and temperature (25°C). There was little effect of irradiance on light CRR, but there was an effect of temperature. Across the seven species light CRR was similar to OCR (oxygen consumption rate in the dark), with the exception of a single species. The outlier species was a coralline alga, and the higher light CRR was probably driven by calcification. CRR could be estimated from OCR, as well as carbon photosynthetic rates from oxygen photosynthetic rates, which suggests that previous studies have probably provided good estimations of gross photosynthesis for seaweeds.