Comparative growth rates and yields of ciliates and heterotrophic dinoflagellates

Growth rates, ingestion rates and grazer yields (grazer volumeproduced/prey volume consumed) were measured for six protozoanspecies (ciliates: Favella sp., Strombidinopsis acuminatum,Uronema sp.; heterotrophic dinoflagellates: Amphidinium sp.,Gymnodinium sp., Noctiluca scintillans) in laboratory batchculture experiments. Comparative growth data indicate that theprymnesiophyte Isochrysis galbana, the prasinophyte Mantoniellasquamata, two cryptophyte species and several autotrophic dinoflagellatespecies were suitable foods for these grazers. When grown onoptimized diets at 13C, maximum ciliate growth rates (range0.77–1.01 day^–1 uniformly exceeded maximum heterotrophicdioflagellate growth rates (range 0.41–0.48 day^–1).A compilation of published data demonstrates that this growthrate difference persists across a range of ciliate and dinoflagellatetaxa and cell sizes. Comparison of volume-specific ingestionrates and yields for the six species studied here showed thatthere was no single explanation for this growth rate disparity.Heterotrophic dinoflagellates exhibited both low ingestion ratesand, in one case, low yields; ciliates were able to achievehigher growth rates via either higher ingestion rates or higheryields, depending on ciliate species. Volume yield increasedover time throughout the exponential growth phase in nearlyall experiments, suggesting variation in response to changingfood concentrations or long-term acclimation to culture conditions.Higher maximum ciliate growth rates mean that these grazershave the potential to exercise tighter control over incipientblooms of their prey than do heterotrophic dinoflagellates.     

Egg production, growth and development of the cyclopoid copepod Oithona similis

Egg production, growth and development rates of Oithona similiswere measured in the laboratory as a function of food concentrationand composition. On an optimum diet, development is isochronaland growth is near exponential. The maximum juvenile growthrate at 15C (0.2 day^–1) is similar to juvenile growthin calanoid copepods. The maximum weight-specific egg productionrate (0.1 day^–1), on the other hand, is substantiallyless than in free-spawning calanoids, but similar to that inegg-carrying calanoids. In the Kattegat, Oithona spp. egg productionis strongly limited by food during summer and controlled bytemperature during winter. The seasonal signal in fecundityand population biomass is much weaker than in the co-occurringfree-spawning calanoid genera, where fecundity and populationbiomass undergo dramatic seasonal viaration. ¹Present address: Instituto Nacional de Investigacin y DesarrolloPesquero, INIDEP, CC 175, Playa Grande, Mar del Plata, Argentina     

Feeding of freshwater filter-feeders at very low food concentrations: Poor evidence for "threshold feeding" and "optimal foraging" in Daphnia longispina and Eudiaptomus gracilis

The feeding behaviour of two freshwater zooplankton species,the calanoid copepod Eudiaptomus gracilis and the cladoceranDaphnia longispina, at extremely low food concentrations hasbeen studied in order to test whether the animals react as predictedby the models of optimal foraging. Three different sized algae were offered in concentrations downto 1µg C/litre at 7°C during winter and 19°C duringsummer. Ingestion rates were measured by use of a flow-through¹⁴C-technique In contrast to the findings of several authors working withmarine copepods a threshold concentration where the animalsceased filtering could be detected only in one experiment byregression analysis (Daphnia, 19°C, food: Stichococcus).A depression of the filtering rate at low concentrations wasobserved only in Daphnia at 19°C with Stichococcus and Scenedesmusas food. It is supposed that in this case the reduction of thefiltering rate is not a direct response of the filtrator, butis caused by exhaustion due to the experimental conditions. As freshwater zooplankton is very rarely exposed to food abundancesas low as in the marine environment, there seems to be no selectionpressure favouring the evolution of an "optimal forager" behaviour. ⁽¹⁾Supported by Deutsche Forschungsgemeinschaft     

Growth and bioluminescence of Noctiluca scintillans on varying algal diets

The effects of food concentration and food quality on the growthrate and bioluminescence potential of the heterotrophic dinoflagellateNoctiluca scintillans were examined. The growth rate of N. scinullansvaried greatly with the phytoplankton food it was provided,and those foods yielding higher growth rates also produced cellsthat were smaller in diameter, but with a greater bioluminescencepotential per unit volume. Small phytoplankton species, includingIsochrysis galbana and an unidentified chrysophyte responsiblefor the Texas brown tide, supported the lowest growth rates.Other small flagellates (Dunaliella tertiolecta) and dinoflagellates(Gyrodinium dorsum, Prorocentrum minimum) supported moderategrowth rates. The highest growth rates were supported by diatomsand prasinophytes. Maximum growth rates (0.5 day^-1) were obtainedwith the diatom Thalassiosira sp. at concentrations     

Canopy Photosynthesis and Crop Growth Rate of Eight Temperate Forage Grasses

The rates of canopy and individual leaf photosynthesis, ratesof growth of shoots and roots, and the extinction coefficientfor light of eight temperate forage grasses were determinedin the field during early autumn. Canopy gross photosynthesiswas calculated as net photosynthesis plus dark respiration adjustedfor temperature using a Q₁₀ = 2. The relationships between canopygross photosynthesis and light intensity were hyperbolic, andthe initial slopes of these curves indicated that light wasbeing utilized efficiently at low light intensities. The initialslope depended on the distribution of light in the canopy andthe quantum efficiency of the individual leaves. The maximumrate of canopy gross photosynthesis reflected the maximum rateof individual leaf photosynthesis. Although the maximum rateof canopy gross photosynthesis was correlated with crop growthrate, there was no significant relationship between daily grossphotosynthesis and crop growth rate. Indeed, daily gross photosynthesisvaried by only 22 per cent, whereas the daily growth of shootsand roots varied by 120 per cent. This poor correlation is influencedby a negative correlation (P < 0.01) between the maximumrate of canopy gross photosynthesis and the initial slope ofthe curve relating canopy gross photosynthesis and light intensity.Difficulties in the interpretation of measurements of dark respirationappeared to confound attempts to relate daily net photosynthesisto crop growth rate, individual leaf photosynthesis, and theextinction coefficient for light.     

Plasticity in filtering screens of Daphnia cucullata×galeata hybrids and parental species at two food concentrations

The coexistence of Daphnia cucullata × galeata hybrids with the parental species D. galeata and D. cucullata was investigated by measuring areas and mesh sizes of filtering structures of these herbivorous zooplankton taxa cultivated at low and high food concentrations. The clearance rates and somatic growth rates were also determined. When reared at low food concentration, all taxa had larger filtering areas. Larger filtering areas also resulted in higher clearance rates. Differences between taxa in both filtering area and clearance rate were caused mainly by interspecific size differences. Hybrids had the largest absolute mesh sizes, and the parental species had smaller mesh sizes. Hybrids also showed heterosis in somatic growth rate at high food concentration. The observed taxon-specific differences in mesh size and somatic growth rate contribute to resource partitioning between the taxa and thus to their successful coexistence in lakes. Received: 1 March 1999 / Accepted: 25 May 1999     

Type-3 functional response in limnetic suspension-feeders,as demonstrated by in situ grazing rates

Field-measured grazing rates (ml/animal/d) of cladocerans (mostly daphniids) and diaptomids were assembled from various published studies and plotted as a function of corresponding phytoplankton concentration (μg l⁻¹ f.w.). Filtering rates of both zooplankton groups initially increased with seston concentration until maximal grazing rates were observed at approximately 4 × 10² and 1 × 10² μg l⁻¹ for cladocerans and copepods, respectively; at higher algal concentrations, filtering rates of both declined as a function of food concentration. The shape of these curves are most consistent with Holling's (1966) Type 3 functional response. We found little support for the Type 3 functional response in published laboratory studies of Daphnia; most investigators report either a Type 1 or Type 2 response. The one study in which the Type 3 response was observed involved experiments where animals were acclimated at low food concentrations for 24 h, whereas those studies associated with response Types 1 or 2 had acclimation periods of only 1 to 3 h. We therefore assembled relevant data from the literature to examine the effect of acclimation period on the feeding rates of Daphnia at low food concentrations. In the absence of any acclimation, animals filtered at extremely low rates. After 2 h of acclimation, however, filtering rates increased 4 to 5-fold but declined again with longer durations; after > 70 h of pre-conditioning, filtering rates were almost as low as they had been with no acclimation. We also found little support for the Type 3 functional response in published studies of copepods. The only study associated with a Type 3 response involved a marine copepod that had been subjected to a starvation period of 48 h; however, an analysis of the effects of acclimation period did not yield conclusive evidence that filtering rates of freshwater copepods (Diaptomus and Eudiaptomus) decrease significantly with acclimation duration. The low filtering rates associated with long acclimation periods in laboratory experiments appears to be a direct result of animals becoming emaciated from prolonged exposure to low food concentrations, a situation which renders them incapable of high filtering rates. This may explain the Type 3 functional response for field cladocerans, since zooplankton in food-limiting situations are constantly exposed to low food concentrations, and would therefore have low body carbon and consequently less energy to filter-feed. We cannot, however, use this to explain the Type 3 response for field diaptomids, since copepods in the laboratory did not appear to lose body carbon even after 72 h of feeding at very low food levels, and there was inconclusive evidence that either Diaptomus or Eudiaptomus decrease their filtering rates with acclimation period. Although Incipient Limiting Concentrations (ILC) for Daphnia ranged from 1 to 8.5 × 10³ μg 1⁻¹, more than half of these fell between 1 and 3 × 10³ μg l⁻¹, bracketing the value of 2.7 × 10² μg l⁻¹ for field cladocerans. There was, however, a great deal of variation in reported maximum ingestion rates (MIR), maximum filtering rates (MFR) and ILC values for Daphnia magna. ILC values from the few laboratory studies of freshwater copepods ranged between 0.5 to 2.8 × 10³ μg 1⁻¹, and was higher than the ILC value of approximately 0.2 × 10³ μg l⁻¹ calculated for field populations of D. minutus. Generally, there was considerable agreement among laboratory studies regarding the shape of grazing-rate and ingestion-rate curves when data were converted to similar units and presented on standardized scales.     

The effects of food deprivation, prey density and volume on clearance rates and ingestion rates of Diacyclops thomasi

The ingestion rates of the copepod, Diacyclops thomasi, on thesoft-bodied rotifer, Synchaeta pectinata, increased 10-fold(0.07–0.77 Synchaeta h^–1) over the range 50–250prey l^–1. The saturating functional response curve appearedsigmoid but was statistically indistinguishable from a parabola.The response curve was more linear and 10 times lower over thesame range of density when Diacyclops was offered Kerarellacochleans, a species having a stiffened lorica. Diacyclops maximizedits ingestion rate on Synchaeta as a function of the availablegut space. Predation effort, measured as clearance rates, waslinked tentatively to changes in swimming speed of Diacyclopsand was a function of hunger level. Diacyclops, which were starvedfor varying periods of time, increased their ingestion rateson Synchaeta up to a maximum (-3.0 h^–1) after 7–10h of food deprivation. The gut passage time of Diacyclops wasestimated to be 7–8 h. Therefore, ingestion rates (andclearance rates) appeared to be strongly correlated to the volumeof food in the gut.     

Day/night differences in the grazing impact of marine copepods

Day/night differences in the removal rate of phytoplankton can occur as a result of increased copepod grazing rates at certain times of the day and diel vertical migration of animals. We conducted shipboard grazing experiments and fine-scale vertical zooplankton sampling to resolve these behaviors. Day/night feeding differences were compared in the center of several warm-core Gulf Stream rings, under conditions of no lateral water mass exchange, in the mesohaline portion of Chesapeake Bay and when following drogues in the Chesapeake Bay plume. Day/night variations in copepod biomass in the surface mixed layer were greater in neritic waters as compared to the open ocean stations. Day/night differences in weight-specific copepod filtration rates varied less than biomass. At the neritic stations copepod grazing was often higher at night, whereas at the oceanic stations day/night grazing rates were similar or daytime grazing rates were highest. The night/day ratio of zooplankton grazing impact on the phytoplankton community (the product of zooplankton biomass and their weight-specific grazing rate) averaged 4.8 in the Chesapeake Bay plume and 1.6 in warm-core Gulf Stream rings. Our results suggest that at lower food levels, there often are less day/night differences in the removal rate of phytoplankton by the copepod community.     

Effects of small-scale turbulence on feeding rate and gross-growth efficiency of three Acartia species (Copepoda: Calanoida)

Experiments designed to test the effects of small-scale turbulenceon the feeding rates and gross-growth efficiency of calanoidcopepods have been performed within a wide range of controlledfood concentrations. Turbulence significantly enhanced feedingrates only at food concentrations lower than the ingestion saturatinglevel. Gross-growth efficiency (the quotient carbon-egg production/carbon-foodingested) of the different Acartia species studied showed differentpatterns of response to turbulence, in agreement with the hydrodynamiccharacteristics of their habitat. Furthermore, experiments conductedon Acartia clausi at two different intensities of turbulenceindicate a shift in the response, with enhancement of feedingat low intensities of turbulence and negative interference athigher intensities.     

Growth Dynamics and Morphology of Annual Shoots, According to their Architectural Position, in Young Cedrus atlantica (Endl.) Manetti ex Carriere (Pinaceae)

The influence of shoot architectural position on growth andbranching pattern of young Cedrus atlantica (Endl.) Manettiex Carrière trees were studied. Extension growth andtype of axillary products (lateral bud, sylleptic short or longshoots) of annual shoots of increasing branching order (mainstem, branches and branchlets) were recorded weekly during the1993 growing season. Annual final shoot length, duration ofextension, and maximum extension rate decreased with increasingbranching order. Sylleptic axillary shoots occurred only onannual shoots of the main stem and branches and were producedwhen extension rate was at its highest. Differences in growthrate and final length of annual shoots, according to their architecturalposition, were related to differences in the total number anddiversity of types of sylleptic axillary shoots produced. Itis suggested that types and numbers of sylleptic axillary shootsproduced are linked with threshold values for both final lengthand extension rate of the parent shoot. Copyright 1999 Annalsof Botany Company Atlas cedar, extension growth, sylleptic branching, tree architecture, morphology.     

Fecal pellet production between molts in a cyclopoid copepod: patterns, individual variability and implications for growth and development

We measured the fecal pellet production rate (FPR) of a cyclopoid copepod (Cyclops cf. sibiricus) individually through the copepodite 5th stage in order to describe the molt-cycle dependent variations in the FPR and the relationship between the FPR and growth and development. Even though individuals of C.cf. sibirucus were kept in the same environmental condition (food quantity, etc.) throughout the development, the FPR was not constant. In general, the FPR was low just after molting into C5, but increased with time and reached the maximum a few days after the molt, and then decreased toward the next molt. More fecal pellets were produced during the earlier part of stage duration time, irrespective of variations in the stage duration time. The total volume of fecal pellets voided through C5 (tCFP) also varied individually with females tending to void more than males. However, neither the weight-specific FPR nor the weight-specific tCFP were different between the sexes. The weight-specific FPR was positively correlated with the individual stage-specific growth rate, suggesting that better growers had a higher ingestion rate, given that the FPR is proportional to the ingestion rate as suggested by other studies.     

A New Generalized Logistic Sigmoid Growth Equation Compared with the Richards Growth Equation

A new sigmoid growth equation is presented for curve-fitting,analysis and simulation of growth curves. Like the logisticgrowth equation, it increases monotonically, with both upperand lower asymptotes. Like the Richards growth equation, itcan have its maximum slope at any value between its minimumand maximum. The new sigmoid equation is unique because it alwaystends towards exponential growth at small sizes or low densities,unlike the Richards equation, which only has this characteristicin part of its range. The new sigmoid equation is thereforeuniquely suitable for circumstances in which growth at smallsizes or low densities is expected to be approximately exponential,and the maximum slope of the growth curve can be at any value.Eleven widely different sigmoid curves were constructed withan exponential form at low values, using an independent algorithm.Sets of 100 variations of sequences of 20 points along eachcurve were created by adding random errors. In general, thenew sigmoid equation fitted the sequences of points as closelyas the original curves that they were generated from. The newsigmoid equation always gave closer fits and more accurate estimatesof the characteristics of the 11 original sigmoid curves thanthe Richards equation. The Richards equation could not estimatethe maximum intrinsic rate of increase (relative growth rate)of several of the curves. Both equations tended to estimatethat points of inflexion were closer to half the maximum sizethan was actually the case; the Richards equation underestimatedasymmetry by more than the new sigmoid equation. When the twoequations were compared by fitting to the example dataset thatwas used in the original presentation of the Richards growthequation, both equations gave good fits. The Richards equationis sometimes suitable for growth processes that may or may notbe close to exponential during initial growth. The new sigmoidis more suitable when initial growth is believed to be generallyclose to exponential, when estimates of maximum relative growthrate are required, or for generic growth simulations.Copyright1999 Annals of Botany Company Asymptote,Cucumis melo,curve-fitting, exponential growth, intrinsic rate of increase, logistic equation, maximum growth rate, model, non-linear least-squares regression, numerical algorithm, point of inflexion, relative growth rate, Richards growth equation, sigmoid growth curve.     

Feeding of dominant zooplankton in Prydz Bay, Antarctica, during austral spring/summer: food availability and species responses

Information on the food and predation of zooplankton species is essential for an improved understanding of zooplankton community dynamics of the Southern Ocean. Feeding of dominant zooplankton species at locations across the neritic, slope and oceanic regions of Prydz Bay, Antarctica, was investigated with incubation experiments during austral spring/summer of 2009/2010 to identify the response of dominant copepods and krill to different feeding environments. Results showed distinct spatial and temporal variations in ingestion and filtration of predominant copepods and krill. In late spring, Calanoides acutus was inactive and the ingestion rate was only 1.30 μgC/ind/day. During early summer, a diatom bloom was in progress at station IS21, showing a peak biomass of Thalassiosira spp. and Fragilaria spp. Daily ingestion rates of Euphausia crystallorophias, C. acutus, Metridia gerlachei and Ctenocalanus citer were relatively high. By contrast, copepod ate predominantly ciliates in slope and oceanic regions where microplankton biomass were lower (<20 μgC/L). During late summer, microplankton of neritic regions, mainly composed of nano-sized Pseudo-nitzschia spp. and ciliates, was less than 10 μgC/L. C. acutus incubated in neritic regions mainly ate ciliates. The total microplankton biomass was lower (<5 μgC/L) and predominated by Pseudo-nitzschia spp. in slope regions north of Fram Bank, and daily ingestion rates of incubated copepods were less than 2.5 μgC/ind. Our results clearly demonstrated that copepods and krill had flexible feeding strategies to cope with temporally and spatially changing food availability in Prydz Bay. Meanwhile, ciliates appeared to represent an important carbon source for zooplankton, especially in regions with lower food concentrations.     

Food-web manipulations influence grazer control of phytoplankton growth rates in Lake Michigan

Stocking piscivorous salmonids in Lake Michigan produced dramaticalterations in food-web structure, including higher numbersof large-bodied zooplankton (especially Daphnia pulicaria),lower summer chlorophyll concentrations and increased watertransparency. Experimental determinations of epilimnetic phytoplanktongrowth rates and of zooplankton grazing rates indicate thatherbivorous zooplankton controlled algal dynamics during thesummer of 1983 because grazers occupied the surface waters throughoutthe day. In 1985, however, both large- and small-bodied Daphniamade approximately equal contributions to total grazer biomass,and all grazers displayed pronounced diel vertical migrations,visiting epilimnetic waters only at night. This prohibited zooplanktonfrom controlling algal dynamics because grazing losses did notexceed phytoplankton growth rates. The changes in zooplanktoncommunity composition and behavior observed in summer 1985 probablyresulted from increased predation by visually orienting planktivorousfish, especially bloater chub (Coregonus hoyi). Effects of food-webmanipulations on phytoplankton dynamics were evident only duringJuly and August. During spring and early summer copepods dominateLake Michigan's zooplankton community. Owing to their smallbody size, copepods are less susceptible to fish predation andexhibit much lower filtering rates than Daphnia. Variabilityin zooplanktivorous fish abundance probably has little effecton phytoplankton dynamics during spring and early summer.     

Relationship between the Growth Rate of a Filament and the Ability to Differentiate a Bud in the Moss, Physcomitrium sphaericum

Protonemata filaments of the moss Physcomitrium sphaericum displaygrowth rates that change remarkably before they differentiatebuds under normal conditions, that is, without cytokinin treatment.Before bud differentiation occurred, the growth rate of thefilament first decreased to a minimum and then increased toa maximum and decreased again. Study of whether the change inthe growth rate is related to bud differentiation showed thatbud differentiation was closely related to the maximum growthrate and the fluctuation range, that is, the difference betweenthe minimum and the succeeding maximum growth rate. The frequencyof bud-differentiated filaments increased as both the maximumgrowth rate and the fluctuation range of growth rate increased.Examination of the relationship between the time of cytokininapplication and the frequency of bud-differentiated filamentsshowed that the cytokinin treatment given at the time when thegrowth rate increased was most effective to induce buds. Therelationships between cytokinin, growth rate of filaments andthe ability to differentiate buds are discussed. (Received September 17, 1985; Accepted March 13, 1986)     

Experimental evaluation of herbivory in the ctenophore Mnemiopsis leidyi relevant to ctenophore-zooplankton-phytoplankton interactions in Narragansett Bay, Rhode Island, USA

Herbivory of Mnemiopsis leidyi and its interactions with phytoplanktonand non-gelatinous zooplankton were examined in small-scalemicrocosm experiments. Clearance rates for M. leidyi incubatedwith phytoplankton were generally negative, but ranged up to4.5 1 ctenophore^–1 day^–1 when the large (80 µmø) diatom Ditylum brightwelli was offered as a food source.These highest ingestion rates would provide Mnemiopsis withonly 21 % of its daily carbon requirements for respiration.Mean shrinkage of M. leidyi was 8.2–51% when incubatedwith phytoplankton. Although M. leidyi neither fed activelyon phytoplankton, nor satisfied its nutritional needs on sucha diet, the chain-forming diatom Skeletonema costatum becameentangled in mucus strands and balls produced by M. leidyi inthe absence of zooplankton. Attachment onto mucus occurred atphytoplankton concentrations commonly observed in NarragansettBay and may be important in the formation of "marine snow" duringsummer M. leidyi pulses; phytoplankton sinking rate and the"package size" available to herbivores would also be affected.The experiments support our previous hypothesis based on fieldobservations in Narragansett Bay that M. leidyi indirectly regulatesphytoplankton abundance there during the summer bloom as a consequenceof predation on zooplankton. The extent to which M. leidyi influencedphytoplankton dynamics in the microcosms was dependent on therelative abundance and physiological state of the three trophiclevels. A food web diagram for M. leidyi is presented.     

Interactions between temperature and light intensity on growth and photosynthesis of the cyanobacterium Oscillatoria agardhii

Oscillatoria agardhii was grown in turbidostat cultures undera 16/8 h light/dark cycle at various combinations of light intensityand temperature. Temperature was found to influence only themaximal growth rate; this relationship was linear over the temperaturerange studied. An equation was derived describing the growthrate (µ) as a continuous function of light intensity andtemperature. The light harvesting pigments chlorophyll a andC-phycocyanin increased in concentration when growth becamelight limited. The regulation patterns observed did not suggestany influence of temperature on their steady state concentrations.The initial slope of the P versus I curves (     

Functional response and food selection of the water flea, Bosmina longispina

The functional response and the feeding behaviour of Bosminalongispina in relation to varying food concentrations were investigatedunder laboratory conditions using radioactively labelled Scenedcsmusacutus and different sized fluorescent latex beads to measureclearance rates and size selectivity. Bosmina longispina reducedfeeding activity at low concentrations of food, having a sigmoidfunctional response. Particles of 5.18 µm were filteredmuch more efficiently than both 1.04- and 0.25-µm particles,and the selectivity of large particles increased with increasingfiltering rates. Scenedesmus acutus was filtered at approximatelythe same rate as 5.18-µm latex beads. The increased selectivityat high filtering rates is explained as an adjustment in thefiltering apparatus to allow for greater water flow. Both thesefeeding responses would tend to reduce energy expenditure insituations when energy intake is low.     

Growth of Near-isogenic Wheat Lines Differing in Development--Spaced Plants

The relationship between phenological development in wheat (TriticumaestivumL.) and growth was studied to determine if the switchfrom a vegetative to a reproductive apex increases plant growthrate. Plant partitioning and relative growth rates during vegetativeand pre-flowering reproductive periods were determined in twosets of near-isogenic lines differing in phenological development.Spaced plants were grown in two photoperiods (15 and 10 h) toincrease the range of development rates. Lines within each isogenicset and photoperiod treatment did not differ in whole plantgrowth rate despite large differences in developmental rate.In addition, the partitioning of biomass between roots and shootswas also similar. The transition of the apex from vegetativeto reproductive mostly affected the partitioning of shoot biomassinto leaf (blades) and stems (rest of the shoot). A longer timeto reach floral initiation was associated with the productionof more, and larger, leaves as well as more tillers. This resultedin large differences in leaf area between isolines. However,at the whole plant level, all lines accumulated biomass at thesame rate with time. The early flowering lines compensated fortheir reduced leaf area by having a higher net assimilationrate and were thereby able to maintain the same relative growthrate as their later flowering counterparts.Copyright 1998 Annalsof Botany Company Development, growth, partitioning,Triticum aestivumL., wheat, isolines.