Surfacing activity and food utilization in a tropical air-breathing fish exposed to different temperatures

Reared in tubular aquaria containing different depths of water (2.5, 5.0, 15.5, 31.0 and 40.0 cm), the obligatory air-breathing fish Ophiocephalus striatus (760 mg; 4.5 cm L) was forced to swim vertically a longer or shorter distance per surfacing. Interaction of temperature (17, 22, 27, 32 and 37°C) and aquarium depth reveals that surfacing frequency of the fish, fed ad libitum on Tilapia muscle, increased with increasing aquarium depth, but the increase was significant only at 27 and 32°C; in the starving series, the frequency was not depth-dependent at any temperature. Owing to the sustained surfacing activity and the consequent fatigue, the test individuals ‘hung’ to the surface for a definite period. Hanging frequency was temperature-dependent, but not a depth-dependent activity either in the starving or feeding series. At any temperature and aquarium depth, the feeding series hung more frequently than the starving series. Hanging duration increased from about 1 hr/day in either series at 17°C to 6 and 18 hr/day in the feeding and starving series at 37°C. At any tested temperature, distance swum by the feeding and starving series was a depth-dependent activity. The feeding series at 32°C exhibited the maximum swimming speed of 2 L/sec for 4.8 hr/day in the 40 cm depth. With increasing temperature and depth, feeding rate increased (from 24 to 225 g cal/g live fish/day); between 17 and 27°C, it was more a temperature-dependent activity. The highest rate (47 g cal / g/day) and efficiency (27%) of conversion were observed at 32°C; whereas the efficiency was depth-dependent, the rate was not. Oxygen uptake was a temperature-dependent activity; aquarium depth played a secondary role. Briefly, O. striatus in deeper aquaria consumed significantly more food and converted lesser, as it surfaced more frequently and swam longer distance, dissipating more energy on metabolism and swimming activity. Hence, culturing O. striatus in shallow waters at the optimum temperature of 32°C will be advantageous. This work was supported by the University Grants Commission's (New Delhi) grant to Dr. T. J. Pandian: Grant No. F. 23-210/75 (Sr II) for which appreciation is expressed.     

Effects of satiation on feeding and swimming behaviour of planktivores

Hunger affects the feeding and swimming behaviour in fish. After 36 h of food deprivation, the feeding and swimming behaviour of Pseudorasbora parva (Cyprinidae) was studied under different prey densities (0.5, 1, 2, 5, 10 and 25 of Daphnia pulex per liter). The initial feeding rates showed marked variations in relation to prey availability. Under high prey densities, the initial feeding rate of fish was higher and subsequently decreased faster, when compared to those feeding under low prey densities. At higher prey densities, two factors were involved: that of higher prey encounter rates and also the attainment of food satiation at a faster rate. Across all prey densities, the feeding rates of fish reached a plateau after satiation. The swimming speed of fish was found to be negatively related to the prey density and a significant change in swimming speed was noted as being directly related to the level of satiation. It was found that the increasing satiation level greatly influenced the handling time and reactive volume of predator, which finally caused reduced feeding rates.     

Foraging behaviour of planktivorous fish in artificial vegetation: the effects on swimming and feeding

In the littoral zones of lakes, aquatic macrophytes produce considerable structural variation that can provide protection to prey communities by hindering predator foraging activity. The swimming and feeding behaviour of a planktivore, Pseudorasbora parva(Cyprinidae) on its prey (Daphnia pulex) was studied in a series of laboratory experiments with varying densities (0, 350, 700, 1400, 2100 and 2800 stems m^–2) of simulated submerged vegetation. Prey availability was varied from 0.5, 1.0, 2.0, 5.0, 10.0 and 25.0 prey l^–1. As the stem density increased, the predator's swimming speed and the number of prey captured decreased relative to feeding in open water. A good relation existed between the number of successful prey captures and swimming speed with the average stem distance to fish body length ratio (D). An abrupt reduction in feeding and swimming was recorded when D was reduced to values less than one.     

Thermal sensitivity of metabolic rates and swimming performance in two latitudinally separated populations of cod, <Emphasis Type="Italic">Gadus morhua</Emphasis> L.

Atlantic cod populations live in a wide thermal range and can differ genetically and physiologically. Thermal sensitivity of metabolic capacity and swimming performance may vary along a latitudinal gradient, to facilitate performance in distinct thermal environments. To evaluate this hypothesis, we compared the thermal sensitivity of performance in two cod stocks from the Northwest Atlantic that differ in their thermal experience: Gulf of St Lawrence (GSL) and Bay of Fundy (BF). We first compared the metabolic, physiological and swimming performance after short-term thermal change to that at the acclimation temperature (7°C) for one stock (GSL), before comparing the performance of the two stocks after short-term thermal change. For cod from GSL, standard metabolism (SMR) increased with temperature, while active metabolism (AMR, measured in the critical swimming tests), EMR (metabolic rate after an exhaustive chase protocol), aerobic scope (AS) and critical swimming speeds (U _crit and U _b–c) were lower at 3°C than 7 or 11°C. In contrast, anaerobic swimming (sprint and burst-coasts in U _crit test) was lower at 11 than 7 or 3°C. Factorial AS (AMR SMR⁻¹) decreased as temperature rose. Time to exhaustion (chase protocol) was not influenced by temperature. The two stocks differed little in the thermal sensitivities of metabolism or swimming. GSL cod had a higher SMR than BF cod despite similar AMR and AS. This led factorial AS to be significantly higher for the southern stock. Despite these metabolic differences, cod from the two stocks did not differ in their U _crit speeds. BF cod were better sprinters at both temperatures. Cod from GSL had a lower aerobic cost of swimming at intermediate speeds than those from BF, particularly at low temperature. Only the activity of cytochrome C oxidase (CCO) in white muscle differed between stocks. No enzymatic correlates were found for swimming capacities, but oxygen consumption was best correlated with CCO activity in the ventricle for both stocks. Overall, the stocks differed in their cost of maintenance, cost of transport and sprint capacity, while maintaining comparable thermal sensitivities.     

Feeding behavior of leaf-like juveniles of the round batfish <Emphasis Type="Italic">Platax orbicularis</Emphasis> (Ephippidae) on reefs of Kuchierabu-jima Island,southern Japan

Feeding behavior of juvenile batfish Platax orbicularis, which presents a mimic state resembling a fallen leaf drifting on the water surface, were surveyed on reefs off Kuchierabu-jima Island, southern Japan. The fish consistently showed drifting swimming patterns on the surface during daylight hours, and fed in a picking–feeding manner on algae covering floating materials or substrates near the water surface. The fish drastically switched to planktivory during the night as they actively hunted free zooplanktonic particles through combined ram and suction feeding manners. Stomach content analysis supported that the fish adopted different feeding tactics, targeting different food items between the two diel periods. Even without an abrupt change of habitat use, due to its mimic state, juveniles of P. orbicularis explore different preys according to their daily activities.     

The Influence of Structural Complexity on Fish–zooplankton Interactions: A Study Using Artificial Submerged Macrophytes

Aquatic macrophytes produce considerable structural variation within the littoral zone and as a result the vegetation provides refuge to prey communities by hindering predator foraging activities. The behavior of planktivorous fish Pseudorasbora parva (Cyprinidae) and their zooplankton prey Daphnia pulex were quantified in a series of laboratory experiments with artificial vegetation at densities of 0, 350, 700, 1400, 2100 and 2800 stemsm^–2. Swimming speeds and foraging rates of the fish were recorded at different prey densities for all stem densities. The foraging efficiency of P. parva decreased significantly with increasing habitat complexity. This decline in feeding efficiency was related to two factors: submerged vegetation impeded swimming behavior and obstructed sight while foraging. This study separated the effects of swimming speed variation and of visual impairment, both due to stems, that led to reduced prey–predator encounters and examined how the reduction of the visual field volume may be predicted using a random encounter model.     

Habitat use and feeding behavior of thirteen species of benthic stream fishes

Synopsis The densities, habitat use, and feeding behaviors of 13 fish species belonging to the benthic invertebrate-feeding guild were studied by snorkeling at three localities in the Little River of eastern Tennessee, U.S.A. Resource partitioning occurred by habitat, feeding behavior and time of activity. Differences were also found at the generic level.Cottus was a nocturnal feeder, whereasPercina andEtheostoma were, for the most part, diurnally active.Percina moved about rapidly and spent most of its time above the bottom. In contrast,Etheostoma varied considerably in the amount of time spent under cover, spent little time above the bottom, and exhibited low levels of swimming activity. Nearly all species sought cover at night, suggesting they may be particularly sensitive to predation at night. Species with small adult sizes (Etheostoma, Cottus andP. evides) were concentrated in shallow water habitats, whereas species with large adult sizes (Percina) were more abundant in deep water habitats. The habitat use data are consistent with the hypothesis that size-selective predation by centrarchid bass may cause smaller fish to avoid deep water areas. Large species should have a lower risk of predation due to their size and behavior.     

Does Acartia clausi (Copepoda: Calanoida) use an area-restricted search foraging strategy to find food?

The swimming behavior of Acartia clausi within small aquaria containing phytoplankton was videotaped, along with unfed controls. From these videos, aspects such as swimming speed, distance traveled, and 2D (horizontal, X, and vertical, Z) headings were then measured from over 75000 X Z positional data in order to quantify their swimming functional response. There was generally no significant difference in swimming behavior between different food levels, although there was a large difference between feeding and unfed controls. Because feeding bouts moved the copepods a short distance at a slow speed, net vertical displacement decreased as feeding-bout frequency increased. Non-feeding individuals showed longer periods of sinking, interspersed with longer-distance, high-speed jumps, which displaced them farther distances than filter-feeding copepods over the same time interval. For both fed and unfed control copepods, a 1D unbiased random walk was adequate to describe their displacement over these very short time periods (10 s). This behavior is consistent with an area-restricted search foraging strategy. The net vertical displacements predicted by combining the experimental data with a 1D random walk model suggest that A. clausi would be retained within even small 10 cm phytoplankton patches long enough to fill their gut. However, copepods outside of phytoplankton patches must rely on other means to find patches of food greater than 50 cm apart. Potentially, knowledge of a copepod's swimming functional response could enable predictions about the typical spatial and temporal patchiness of its food in situ.     

Effects of temperature,swimming speed and body mass on standard and active metabolic rate in vendace (<Emphasis Type="Italic">Coregonus albula</Emphasis>)

This study gives an integrated analysis of the effects of temperature, swimming speed and body mass on standard metabolism and aerobic swimming performance in vendace (Coregonus albula (L.)). The metabolic rate was investigated at 4, 8 and 15°C using one flow-through respirometer and two intermittent-flow swim tunnels. We found that the standard metabolic rate (SMR), which increased significantly with temperature, accounted for up to 2/3 of the total swimming costs at optimum speed (U _opt), although mean U _opt was high, ranging from 2.0 to 2.8 body lengths per second. Net swimming costs increased with swimming speed, but showed no clear trend with temperature. The influence of body mass on the metabolic rate varied with temperature and activity level resulting in scaling exponents (b) of 0.71–0.94. A multivariate regression analysis was performed to integrate the effects of temperature, speed and mass (AMR = 0.82M ^0.93 exp(0.07T) + 0.43M ^0.93 U ^2.03). The regression analysis showed that temperature affects standard but not net active metabolic costs in this species. Further, we conclude that a low speed exponent, high optimum speeds and high ratios of standard to activity costs suggest a remarkably efficient swimming performance in vendace.     

Food and feeding of aquatic larvae of the midge chaoborus flavicans (meigen) (diptera : chaoboridae) in the laboratory

Laboratory observations on the feeding of fourth instar larvae of Chaoborus flavicans (Meigen) show that features relating to the prey, such as swimming behaviour, size and shape, play a major role in determining the kind of food eaten by the predator.In tests with two interacting limnetic cyclopoid copepods, Cyclops abyssorum out-competed Mesocyclops leuckarti when Chaoborus was absent but did not do so when Chaoborus was present.     

Distribution of behaviours and species interactions within home range contours in five Caribbean reef fish species (Family Labridae)

Synopsis This study investigated the distribution of behaviours and species interactions within home range contours in five Caribbean labrid species: Halichoeres bivittatus, H. garnoti, H. maculipinna, H. poeyi, and Thalassoma bifasciatum. For this study, contours were defined as: (a) 30%—the core use area, (b) 30–75%—the intermediate activity area, and (c) 75–95%—the peripheral activity area. Behaviours analyzed for this study included: (i) feeding (=biting the substrate or chewing), (ii) chased by pomacentrids, (iii) swimming alone, (iv) swimming with other fishes, and (v) all activities with other fishes. Fifty-nine percent of Halichoeres bivittatus observed showed a higher frequency than expected being chased by pomacentrids in the peripheral region of their home ranges. Halichoeres garnoti showed a lower frequency than expected swimming with other individuals in their core use area, and 64% of the individuals observed showed a higher frequency than expected being chased by pomacentrids in the peripheral region. In general, H. maculipinna exhibited a random distribution of behaviours throughout their home range areas, with a non-significant trend for more agonistic interactions with pomacentrids in peripheral regions. Halichoeres poeyi and T. bifasciatum showed higher frequencies than expected being chased by pomacentrids in the peripheral regions. Overall, the non-random distribution of agonistic interactions with pomacentrids throughout home range areas suggests that the presence or prior residence of territorial pomacentrids on coral reefs may modify the post-settlement selection of home range areas by these labrid species.     

Swimming Behaviour of <Emphasis Type="Italic">Chironomus acerbiphilus</Emphasis> Larvae in Lake Katanuma

We conducted a seasonal survey of the swimming behaviour of Chironomus acerbiphilus larvae in volcanic Lake Katanuma from April 1998 to December 2001. Swimming C. acerbiphilus density was much higher than other chironomid species in lakes. All C. acerbiphilus larvae (1st through 4th instars) swam, but the earlier instars (especially the 1st) had the greatest densities and fluctuations. First instars were never found in the benthic population. This result indicates that the 1st-instar larvae are planktonic. Low water temperature (below about 10 °C) resulted in the seasonal disappearance of swimming chironomid larvae. Chemical factors – oxygen depletion or presence of hydrogen sulfide – also restricted the distribution of swimming and benthic larvae. Larvae were distributed only in the oxygen-rich part of the lake bottom and swam only in the oxygen-rich layer of the water column. The density of older swimming C. acerbiphilus (3rd and 4th instars) tended to increase with increasing benthic larval densities. The chemical stress of oxygen depletion or presence of hydrogen sulfide during holomixis within and after the stratification period leads to conspicuous swimming behaviour of benthic C. acerbiphilus larvae. Almost all C. acerbiphilus larvae died on this occasion.     

Why do electric fishes swim backwards? An hypothesis based on gymnotiform foraging behavior interpreted through sensory constraints

Synopsis Fishes producing high-frequency wavelike electrical discharges maintain a relatively rigid body axis and swim forwards and backwards with equal ease. Using stop-action videotape filming we have observed the gymnotiform Apteronotus albifrons feeding on zooplankton and oligochaete annelids. Here it is reported that reverse swimming is characteristic of two foraging behaviors: searching for prey and assessing it. In assessing a potential prey item, fish typically scan it from tail to head by swimming backwards, then ingest it after a short forward lunge. A scan in the opposite direction-from head to tail by forward swimming-would have the prey located near the tail and out of position for the final lunge. Food choice experiments indicate that these electrosensing fish feed equally well, and take larger rather than smaller zooplankton, under light and dark conditions. Furthermore, electric fish take normal (light) colored and darkened prey (Daphnia) in a 50: 50 ratio under both dark and light conditions. These results are consistent with the interpretation that electrosensory cues are being used to detect zooplankton and other prey. Together, our observations support Lissmann's (1958, 1974) and Lissmann & Machin's (1958) assertion that backwards swimming is a component of a locomotory pattern guided by the constraints produced by an active electrical sense.     

The swimming and feeding behavior of Mesocyclops

The swimming and feeding behaviors of Mesocyclops are described from a review of the literature and personal observations. Mesocyclops exhibits considerable behavioral flexibility in response to environmental stimuli. Mesocyclops edax exhibits an increase in horizontal looping behavior at high prey densities, and performs a tight vertical looping behavior in response to the loss of captured prey. Ingestion rates by Mesocyclops are a complex function of prey density, morphology, and behavior in addition to prey size. Vertebrate predators induce a rapid escape response in Mesocyclops and may be responsible at least in part for their extensive diel vertical migrations. The complex behavioral patterns of Mesocyclops suggest that its distribution and abundance in nature will be distinctly nonrandom and influenced as much by its own behavioral responses as by other external physical factors such as water circulation patterns.     

Functional analysis of a large non-conserved region of FlgK (HAP1) from <Emphasis Type="Italic">Rhodobacter sphaeroides</Emphasis>

The single subpolar flagellum of Rhodobacter sphaeroides shows an enlarged hook-filament junction. One of the two proteins that compose this section of the filament is HAP1 _Rs (FlgK _Rs ) it contains a central non-conserved region of 860 amino acids that makes this protein about three times larger than its homologue in Salmonella enterica serovar Typhimurium. We investigated the role of this central portion of the unusually large HAP1 protein of R. sphaeroides by monitoring the effects of serial deletions in flgK _Rs , the gene encoding HAP1 _Rs , on swimming and swarming. Two deletion mutants did not assemble functional flagella, two were paralyzed and five exhibited reduced free-swimming speeds. Some mutants produced unusual swarming patterns on soft agar without or with Ficoll 400. A segment of approximately 200-aa of the central region of HAP1 _Rs that aligns with the variable region of the flagellin sequence from other γ- and β-proteobacteria was also found. Therefore, it is possible that the origin of this large central domain of HAP1 _Rs could be associated with an event of horizontal transfer and subsequent duplications and/or insertions.     

The components of feeding behavior in rotifers

Feeding behavior of a rotifer can be broken into two classes of activities: the rate of successful search and the handling process. The former consists of the following components: Perceptual field (in planktonic rotifers the area of the corona), swimming rate, and attack rate. The second class consists of capture rate, handling time, rejection rate, ingestion, digestion, and assimilation. All evidence indicates that the perceptual field cannot be varied by the rotifer. Swimming rate is variable and under rotifer control, but does not appear to vary with degree of starvation. Attack rate is also under control of the rotifer, at least in the genus Asplanchna. Capture rates vary with the species of food item from zero to 100%. Handling times are longer than one would expect, as are rejection times. Digestion and assimilation appear to vary inversely with rate of ingestion. There is some suggestion in the literature that feeding behavior on very small particles differs from that on larger ones.     

Modification of the swimming behaviour of Brachionus calyciflorus (Pallas) according to food environment and individual nutritive state

The swimming behaviour of orthoclonal Brachionus calyciflorus (Pallas) females was studied with the aid of a system of automated trajectometry. The swimming was observed in response to the food environment and the individual's nutritive state. Eighteen hours before testing, neonates were placed either in a suspension of Chlorella, or in a freshwater medium. For each nutritive state, the test consisted of an analysis of the swimming behaviour of B. calyciflorus in 5 different environmental food conditions: in an algal suspension, in an algal solution (obtained from filtration of the algal suspension), in a freshwater medium, in the same freshwater medium with a suspension of polystyrene beads, and in the algal solution with the same suspension of beads. Three variables were used to describe the swimming path: linear speed (mm s^–1), angular speed (degrees s^–1), and mean angle (degrees). The results showed a modification in the swimming pattern dependent on both food environment and nutritive state of females. The nature of stimulations inducing these behavioural modifications, and their interactions with the individual nutritive state are briefly discussed.     

Studying <Emphasis Type="Italic">Daphnia</Emphasis> feeding behavior as a black box: a novel electrochemical approach

We present a novel approach for examining the complex feeding behavior of a filter feeder at a previously unexploited scale. A Daphnia lives in a viscous environment and thus creates a feeding current with a distinct laminar inflow and a repetitive pulsed outflow. We propose that by treating the feeding apparatus as a black box, and using the pulsed outflow current as a surrogate to the inside working of the apparatus, we can calculate feeding rate in near real time. The structure of the outflow is interpreted as a direct representation of the organism’s response to its environment. Therefore, we examine how the work performed by an organism’s feeding apparatus is altered according to environmental factors and metabolic demands. Our approach is an integration of optical (Schlieren system) and electrochemical (chronoamperometry) techniques that allow for real time visualization and temporal analysis of flow systems, respectively. As electrochemistry requires a tracer chemical, we employed low dopamine concentrations (≤ 1mM), and tested the effect of dopamine on the heart rate and swimming of Daphnia. It appears that dopamine free in solution at concentrations below 10 mM has no adverse effects on the organism, and all observed differences in Daphnia feeding behavior were due to environmental or metabolic factors. The feeding nature of daphnids in the presence or absence of food, and differences between the sexes is reported. Our results indicate that in the absence of food a Daphnia has a strict and repetitive feeding behavior with short delays between pumping actions. However, in the presence of food this behavior becomes complex, with increased delays between pumps, perhaps designed to maximize feeding efficiency. Our observations demonstrate that males have a higher appendage beat frequency than females under identical conditions. We hypothesize that the difference may be dictated by metabolic demand, as a male spends more time actively seeking a mate. The application of electrochemistry to the study of Daphnia feeding behavior is an improvement over current methods for its near real time quantification of behavioral response, its versatile application under varying environmental conditions and its extreme sensitivity to changes in the organism’s feeding behavior. This technique is a valuable addition to the current tools available for studying Daphnia feeding behavior and will allow us to learn more about the interactions of an organism with its environment. Guest editor: Piet Spaak Cladocera: Proceedings of the 7th International Symposium on Cladocera     

Performance of a generalist grasshopper on a C<Subscript>3</Subscript> and a C<Subscript>4</Subscript> grass: compensation for the effects of elevated CO<Subscript>2</Subscript> on plant nutritional quality

The increasing CO₂ concentration in Earths atmosphere is expected to cause a greater decline in the nutritional quality of C₃ than C₄ plants. As a compensatory response, herbivorous insects may increase their feeding disproportionately on C₃ plants. These hypotheses were tested by growing the grasses Lolium multiflorum C₃) and Bouteloua curtipendula C₄) at ambient (370 ppm) and elevated (740 ppm) CO₂ levels in open top chambers in the field, and comparing the growth and digestive efficiencies of the generalist grasshopper Melanoplus sanguinipes on each of the four plant × CO₂ treatment combinations. As expected, the nutritional quality of the C₃ grass declined to a greater extent than did that of the C₄ grass at elevated CO₂; protein levels declined in the C₃ grass, while levels of carbohydrates (sugar, fructan and starch) increased. However, M. sanguinipes did not significantly increase its consumption rate to compensate for the lower nutritional quality of the C₃ grass grown under elevated CO₂. Instead, these grasshoppers appear to use post-ingestive mechanisms to maintain their growth rates on the C₃ grass under elevated CO₂. Consumption rates of the C₃ and C₄ grasses were also similar, demonstrating a lack of compensatory feeding on the C₄ grass. We also examined the relative efficiencies of nutrient utilization from a C₃ and C₄ grass by M. sanguinipes to test the basis for the C₄ plant avoidance hypothesis. Contrary to this hypothesis, neither protein nor sugar was digested with a lower efficiency from the C₄ grass than from the C₃ grass. A novel finding of this study is that fructan, a potentially large carbohydrate source in C₃ grasses, is utilized by grasshoppers. Based on the higher nutrient levels in the C₃ grass and the better growth performance of M. sanguinipes on this grass at both CO₂ levels, we conclude that C₃ grasses are likely to remain better host plants than C₄ grasses in future CO₂ conditions.     

Response of heart rate and cloacal ventilation in the bimodally respiring freshwater turtle, Rheodytes leukops, to experimental changes in aquatic PO2

Changes in heart rate (f _H) and cloacal ventilation frequency (f _C) were investigated in the Fitzroy turtle, Rheodytes leukops, under normoxic (17.85 kPa) and hypoxic (3.79 kPa) conditions at 25°C. Given R. leukops’ high reliance on aquatic respiration via the cloacal bursae, the objective of this study was to examine the effect of varying aquatic PO₂ levels upon the expression of a bradycardia in a freely diving, bimodally respiring turtle. In normoxia, mean diving f _H and f _C for R. leukops remained constant with increasing submergence length, indicating that a bradycardia failed to develop during extended dives of up to 3 days. Alternatively, exposure to aquatic hypoxia resulted in the expression of a bradycardia as recorded by a decreasing mean diving f _H with increasing dive duration. The observed bradycardia is attributed to a hypoxic-induced metabolic depression, possibly facilitated by a concurrent decrease in f _C. Results suggest that R. leukops alters its strategy from aquatic O₂ extraction via cloacal respiration in normoxia to O₂ conservation when exposed to aquatic hypoxia for the purpose of extending dive duration. Upon surfacing, a significant tachycardia was observed for R. leukops regardless of aquatic PO₂, presumably functioning to rapidly equilibrate blood and tissue gas tensions with alveolar gas to reduce surfacing duration.