FIELD STUDIES ON THE GIANT KELP NEREOCYSTIS1,2

An intertidal population of Nereocystis luetkeana was investigated in the field, and techniques for measuring sporophyte growth were developed. Primary regions of growth in plants prior to reproductive maturity were located in the bases of the blades, upper stipe, and holdfast. After reproductive areas on blades appeared, growth was confined to 2 major regions: the blade bases and the junction between lower stipe and holdfast where new holdfast haptera were produced. Complete deblading halted growth at whatever stage the plants had attained at the time of the operation but did not kill the plants. Partial deblading resulted in low but measurable growth rates in the blade bases. The data presented suggest that the function of the blades is twofold: they bear reproductive structures in the mature plants and. they may provide, materials used, in rapidly growing regions.     

Attachment strength differs amongst life‐history stages of an intertidal,isomorphic red alga

Complex haploid‐diploid life cycles amongst marine organisms may be maintained by ecological differences in life‐history phases. For red algal species within the Gigartinaceae, such differences may be driven, in part, by different cell wall composition and resultant biomechanical strengths of haploid and diploid phases. A field experiment tested the attachment strengths of gametophytes and tetrasporophytes of the isomorphic red alga, Chondrus verrucosus (with comparisons of fertile and vegetative fronds, with and without natural tissue damage across three wave‐exposed sites). Seventy‐nine percent of all fronds broke at the stipe‐holdfast junction. There were significant differences in attachment strength (break force and break stress), but not gross morphology (frond length, number of branch axes, wet weight and cross‐sectional area of fronds that dislodged at the stipe‐holdfast junction) of life‐history phases, with tetrasporophytes exhibiting weaker tissue strength and attachment, and therefore greater susceptibility to dislodgement by waves. However, fertility and tissue damage did not consistently influence dislodgement in pull‐to‐break tests simulating the effects of single waves. The ecological and evolutionary consequences of greater susceptibility to dislodgement of tetrasporophytes (relative to gametophytes) warrant further investigation.     

Endoplasmic filaments generate the motive force for rotational streaming in Nitella

The streaming endoplasm of characean cells has been shown to contain previously unreported endoplasmic filaments along which bending waves are observed under the light microscope using special techniques. The bending waves are similar to those propagated along sperm tails causing propulsion of sperm. In Nitella there is reason to believe that nearly all of the filaments are anchored in the cortex and that their beating propels the endoplasm in which they are suspended. This hypothesis is supported by calculations in which typical and average wave parameters have been inserted into the classical hydrodynamic equations derived for sperm tail bending waves. These calculations come within an order of magnitude of predicting the velocity of streaming and they show that waves of the character described, propagated along an estimated 52 m of endoplasmic filaments per cell, must generate a total motive force per cell within less than an order of magnitude of the forces measured experimentally by others. If we assume that undulating filaments produce the force driving the endoplasm, then the method described for measuring the motive force could lead to a lower than actual value for the motive force, since both centrifugation and vacuolar perfusion would reverse the orientation of some filaments. Observations of the initiation of particle translation in association with the filaments suggest that particle transport and wave propagation, which occur at the same velocity, may both be dependent on the same process. The possibility that some form of contractility provides the motive force for filament flection and particle transport is discussed.     

Metabolic consequences of living in a wave-swept environment: Effects of simulated wave forces on oxygen consumption, heart rate, and activity of the shell adductor muscle of the abalone Haliotis iris

Animals in wave-exposed habitats must constantly contend with the hydrodynamic forces of lift and drag. In this study, we investigated aspects of the metabolic response of Haliotis iris to simulated wave forces varying in magnitude up to 9.6 N applied to the shell at 69° to horizontal, alternately from anterior and posterior directions, with a period of 10s. Shell adductor muscle activity (electromyogram, EMG), heart rate, and oxygen consumption were monitored during force application and during extended recovery. EMG spiking was absent at zero force, but increased markedly with increasing force, in synchrony with the wave cycle. In contrast, heart rate was unaffected by wave forces and varied by only 5% over the whole range of applied forces. During force application, oxygen consumption increased by 10-25% above resting rates and remained elevated throughout a 5-hour recovery period, indicating a switch to anaerobic metabolism. It is concluded that living in a wave-swept environment is metabolically costly for abalone although this may be compensated by improved food availability and more efficient ventilation induced by external flow.     

Two decades of biomonitoring polar bear health in Greenland: a review

Background

Different animal models are used as fracture models in orthopaedic research prior to implant use in humans, although biomechanical forces can differ to a great extend between species due to variable anatomic conditions, particularly with regard to the gait. The rabbit is an often used fracture model, but biomechanical data are very rare. The objective of the present study was to measure axial forces, bending moments, and bending axis directly in the rabbit tibia in vivo. The following hypothesis was tested: Axial forces and bending moments in the mid-diaphysis of rabbit tibia differ from other experimental animals or indirectly calculated data.

Methods

A minifixateur system with 4 force sensors was developed and attached to rabbit tibia (n = 4), which were subsequently ostectomised. Axial forces, bending moments and bending angles were calculated telemetrically during weight bearing in motion between 6 and 42 days post operation.

Results

Highest single values were 201% body weight [% bw] for axial forces and 409% bw cm for bending moments. Whereas there was a continous decrease in axial forces over time after day 10 (P = 0.03 on day 15), a decrease in bending moments was inconsistent (P = 0.03 on day 27). High values for bending moments were frequently, but not consistently, associated with high values for axial forces.

Conclusion

Axial forces in rabbit tibia exceeded axial forces in sheep, and differed from indirectly calculated data. The rabbit is an appropriate fracture model because axial loads and bending moments in rabbit tibia were more closely to human conditions than in sheep tibia as an animal model.     

Studies on nitrate reductase activity in Laminariadigitata (Huds.) Lamour. I. Longitudinal and transverse profiles of nitrate reductase activity within the thallus

The distribution of the enzyme nitrate reductase (NR) within the thallus of the brown alga Laminaria digitata (Huds.) Lamour is described for plants sampled from the east coast of Scotland in May and June when growth rates are at a maximum. Highest NR activities (≈ 0.2 μmol NO₃^? reduced·g^?1 wet wt·h^?1) occurred in the mature blade. NR activities declined towards the basal meristematic region of the blade. Activities in the stipe and holdfast were also low, being between 0.05 and 0.07 μmol NO₃^? reduced·g^?1 wet wt·h^?1. The activities of the enzyme glutamine synthetase (GS), which is important in the assimilation of NH₄⁺, showed a similar distribution within the blade to those of NR.The transverse profile of NR activity in the stipe exhibited a decline from the outer to the inner tissues. Maximum activities (0.13 μmol NO₃^? reduced·g^?1 wet wt·h^?1) occurred in the meristoderm, while those of the cortex and medulla were 0.04 and 0.01 μmol NO₃^? reduced·g^?1 wet wt·h^?1 respectively.These data indicate that most of the NO₃^? assimilation occurs in the mature blade rather than in the meristematic tissue where there is a high nitrogen demand for growth. The data are consistent with the maintenance of meristematic growth by the internal transport of organic nitrogen from the mature blade.     

Relation between size and age of holdfasts of Ecklonia stolonifera Okamura (Laminariales,Phaeophyta) in northern Honshu,Japan

Ecklonia stolonifera is distributed along the coast facing the Sea of Japan. The size of various parts of the shoot (blade length and width and stipe length and diameter) and the age were determined at Ooma, Aomori Prefecture. The smaller the holdfast, the higher the percentage of one-year-old shoots. Holdfasts 10 cm in diameter seemed to be three years old, whereas holdfasts 40 cm in diameter seemed to be five or more years old. Zoosporangial sori were observed on blades three or more years old. Ecklonia stolonifera holdfast diameter expands only vegetatively by stoloniferous rhizoids. Zoospores, formed on shoots three or more years old, serve for the formation of new populations.     

Effect of pre-incubation irradiance on survival of cryopreserved gametophytes of Undaria pinnatifida (Phaeophyta) and morphology of sporophytes formed from the gametophytes

Gametophyte strains originating from indigenous sporophytes of Undaria pinnatifida (Harvey) Suringar in Iwate Prefecture, Northeast Japan, were maintained for 9–10 months at 45 μmol photons m⁻² s⁻¹. Before cryopreservation in liquid nitrogen for more than 12 h (1–14 days) using a two-step cooling method with a mixture of cryoprotectants (10% l-proline and 10% glycerol), these were pre-incubated for 2, 4 and 8 months at 15 μmol photons m⁻² s⁻¹. After 1 week of thawing, no surviving gametophytes were detected in the strains without pre-incubation, but both the female and male gametophytes, pre-incubated for more than 4 months, showed high survival rates (43–60% for females and 64–100% for males). This revealed the induction of freezing tolerance by incubation at low irradiance. Thereafter, sporophytes derived from cryopreserved gametophytes and subcultured gametophytes, stored under pre-incubation conditions, were formed from the strain, and a morphological comparison was conducted with 10 characters (stipe length, stipe wet weight, blade length, blade wet weight, blade width, incision depth, blade thickness, sporophyll length, sporophyll wet weight, and sporophyll width). The morphology of the sporophytes formed from the cryopreserved gametophytes corresponded well with that of the subcultured gametophytes from the same strain. The results suggest that the cryopreservation method is applicable for preserving culture stocks of U. pinnatifida to be used in mariculture.     

Hydrodynamic calculations on the movements of cilia and flagella I. Paramecium

Calculations of the velocity profile, force, moment and bending moment using a theoretical model are carried out for the three-dimensional “conical-helical” beat of a cilium of Paramecium multimicro-nucleatum. The mean velocity profile obtained by numerical computation is found to be twisted in form: it is inclined at a slight angle to the effective stroke at the top of the cilia sublayer, but twists around with the recovery stroke in the lower part of the sublayer. The force and moment are larger during the fast effective stroke, but over a complete cycle their contributions are approximately zero. Calculations on the bending moments reveals that they are larger during the recovery stage of the beating cycle.     

Red algae respond to waves: morphological and mechanical variation in Mastocarpus papillatus along a gradient of force

Intertidal algae are exposed to the potentially severe drag forces generated by crashing waves, and several species of brown algae respond, in part, by varying the strength of their stipe material. In contrast, previous measurements have suggested that the material strength of red algae is constant across wave exposures. Here, we reexamine the responses to drag of the intertidal red alga Mastocarpus papillatus Kutzing. By measuring individuals at multiple sites along a known force gradient, we discern responses overlooked by previous methods, which compared groups of individuals between "exposed" and "protected" sites. This improved resolution reveals that material strength and stipe cross-sectional area are both positively correlated with drag, suggesting that individual blades or populations can adjust either or both of these parameters in response to their mechanical environment. The combined effect of this variation is a stipe breaking force that is positively correlated with locally imposed drag. Owing to this response to drag, the estimated wave-imposed limit to thallus size in M. papillatus is larger than previously predicted and larger than sizes observed in the field, indicating that factors other than wave force alone constrain the size of this alga on wave-swept shores.     

MORPHOLOGY AND MOLECULAR PHYLOGENY OF AUREOPHYCUS ALEUTICUS GEN. ET SP. NOV. (LAMINARIALES,PHAEOPHYCEAE) FROM THE ALEUTIAN ISLANDS1

A previously unknown species of kelp was collected on Kagamil Island, Aleutian Islands. The species can be easily distinguished from any known laminarialean alga: the erect sporophytic thallus is composed of a thin lanceolate blade attaining ～2 m in height and ～0.50 m in width, without midrib, and the edge of the blade at the transition zone is thickened to form a V‐shape; the stipe is solid and flattened, slightly translucent, attaining ～1 m in length; the holdfast is semidiscoidal and up to 0.15 m in diameter. Anatomically, the blade has the typical trumpet‐shaped hyphae characteristic of the Chordaceae and derived foliose laminarialean species (i.e., Alariaceae/Laminariaceae/Lessoniaceae). No hair pits or mucilaginous structures were observed on the blade or stipe. No fertile sporophytes were collected, but abundant juvenile sporophytes were observed in the field. In the molecular phylogenetic analyses using chloroplast rbcL gene, nuclear ITS1‐5.8S‐ITS2 rDNA, and mitochondria nad6 DNA sequences, the new species (Aureophycus aleuticus gen. et sp. nov.) showed a closer relationship with Alariaceae of conventional taxonomy, or the “Group 1” clade of Lane et al. (2006) including Alaria and related taxa than with other groups, although the species was not clearly included in the group. Aureophycus may be a key species in elucidating the evolution of the Alariaceae within the Laminariales. Because of the lack of information on reproductive organs and insufficient resolution of the molecular analyses, we refrain from assigning the new species to a family, but we place the new species in a new genus in the Laminariales.     

CARBON ALLOCATION IN MACROCYSTIS PYRIFERA (PHAEOPHYTA): INTRINSIC VARIABILITY IN PHOTOSYNTHESIS AND RESPIRATION1

Measurements of net photosynthesis (PS, O₂ evolution), dark respiration (R, O₂ consumption), and light and dark carbon fixation (¹⁴C) were conducted on whole blades, isolated blade discs, sporophylls, apical scimitars and representative portions of stipe and holdfast of the giant kelp Macrocystis pyrifera L.C. Ag. On a dry weight basis, highest net PS rates were observed in apical scimitar segments and whole blades (3.81 and 3.07 mgC · g dry wt^?1· h^?1, respectively), followed by sporophylls (1.42 mgC·g dry wt^?1· h^?1) and stipe segments (0.15 mgC·g dry wt^?1· h^?1). No PS capacity was observed in holdfast material. Respiration rates showed similar ranking ranging from 1.22 mgC·g dry wt^?1·h^?1 for apical scimitar to 0.18–0.22 mgC·g dry wt^?1· h^?1 for holdfast material. Considerable within blade variability in both PS and R was also found. Steepest PS and R gradients on both an areal and weight basis were found within immature blades followed by senescent and mature blade material. Highest net PS rates were associated with the blade tips ranging from 3.08 (mature blades) to 10.3 mgC·dry wt^?1·h^?1 (immature blades). Highest rates of R generally occurred towards the basal portions of blades and ranged from 1.03–1.80 mgC·g dry wt^?1·h^?1 for immature blades. The variability within and between blades was high, with coefficients of variation approaching 50%. The observed patterns can be related to the decreasing proportionment of photosynthetic tissue and increasing proportionment of structural tissue as occurs from the blade tip to the blade base. Rates of light carbon fixation (LCF) revealed longitudinal profiles similar to oxygen measurements for the different blade types, with the absolute rates being slightly lower. Patterns of dark carbon fixation (DCF) were less easily interpreted. Highest rates of DCF (0.04–0.06 mgC·g dry wt^?1·h^?1) occurred at the basal portions of immature and senescent blades. Longitudinal profiles of total chlorophyll (a + c) on both an areal and weight basis were very similar to the profiles of PS. Normalized to chlorophyll a, PS displayed an unusual longitudinal profile in immature tissue; however, such profiles for mature and senescent tissues were similar to those for PS on an areal basis. It was demonstrated that it is difficult, if not impossible, to select single tissue discs that are representative of whole blades. The metabolic longitudinal profiles reveal a characteristic developmental pattern; the previous working definitions of immature, mature, and senescent blades, based on morphology and frond position thus have a physiological basis.     

Biomechanical properties and holdfast morphology of coenocytic algae (Halimedales, Chlorophyta) in Bocas del Toro, Panama

For attached marine organisms, specific biomechanical properties may result in detachment or in tissue loss, when sufficient tensile force is applied. Algae experience such forces through water movement, which may thus act to limit size, abundance, and species composition, of populations of algae.Coenocytic construction is uncommon in the algae, but it occurs relatively more frequently in green algae found in shallow subtidal sediments associated with coral reefs, e.g., at our study site of Isla Colon, Bocas del Toro, Panama. We studied the biomechanical properties of some tropical coenocytic algae (Udotea flabellum (Ellis et Solander) Lamouroux, Penicillus capitatus Lamarck, P. pyriformis A. and E.S. Gepp, and Halimeda gracilis Harvey) anchored in sediments. We compare our results with published data on other coenocytic algae, as well as with multicellular algae. Our results show that properties of sand-dwelling coenocytes, such as mean force to dislodge (4.9-12.7 N), mean force to break (6.6-22.1 N), and mean strength (1.0-7.0 MN m^− 2), are all within the range reported for temperate, multicellular, algae. In contrast, the coenocytes differed markedly from the temperate non-coenocytes in the consequences of applied tensile force: coenocytes were removed whole, while most temperate algae attached to rocks break within the thallus. Some multicellular algae can regrow from tissue left on the substratum; three of the four coenocytic species we examined had rhizoids connecting closely adjacent (0.1-0.15 m) individuals, and these rhizoids may serve to regrow a new individual. While our experiments indicated that sufficient tensile force results in dislodgment, calculations using the experimentally determined variables led us to conclude that water velocities sufficient to dislodge individuals are unlikely to occur. Since dislodgment is usually fatal for algae, the role of the holdfast is a critical one. All of the species we investigated had similar holdfast morphology, a mass of rhizoids which entrained sand, the entire unit forming a hemispherical to cylindrical mass. Despite the consistency in holdfast form, and the initial prediction that this was an optimal form for anchoring these algae, our data suggest that this is not the case.     

CONTRASTING PATTERNS OF ALLOMETRY AND REALIZED PLASTICITY IN THE SISTER SPECIES MAZZAELLA SPLENDENS AND MAZZAELLA LINEARIS (RHODOPOHYTA)1

Phenotypic differences between the low wave exposure Mazzaella splendens (Setchell et Gardner) Hommersand and the high exposure Mazzaella linearis (Setchell et Gardner) Fredericq could be due to plasticity or genetic differentiation. Common gardens were used to assess their levels of plasticity after describing allometric relationships. As thalli lengthened, stipe development for M. splendens almost ceased even though blades continued to expand, but M. linearis formed a larger stipe before developing a blade that continued to stay narrow at longer thallus lengths. Common gardens demonstrated that M. splendens regrown in the site of M. linearis produced a wider blade than M. splendens regrown in its natural low energy site and that M. linearis regrown in low wave energy either could not form a wider blade or became narrower than thalli from its high energy site. Tetrasporophytes of M. splendens produced a longer and thicker stipe in the high energy site, but the larger M. linearis stipe was not realized because its wider blades made it vulnerable to hydrodynamic removal. Mazzaella splendens therefore had low survivorship in the high wave energy site, and survivors were not long enough to reproduce. Survivorship and reproduction of M. linearis was similar in both environments. Some of the M. splendens and M. linearis characters are plastic, but this plasticity was insufficient for convergence of phenotypes, and blade width plasticity was maladaptive at least for M. splendens. Developmental systems producing the stipe and blade phenotypes of each species have undergone genetic differentiation.     

Growth and reproductive patterns of <Emphasis Type="Italic">Undaria pinnatifida</Emphasis> sporophytes in a cultivation farm in Busan,Korea

Monthly growth and reproduction of Undaria pinnatifida sporophytes were examined over a period of 5 months in a cultivation farm in Korea. A total of 11 characters of Undaria were measured to determine a reliable morphological character representing its growth and reproduction. Plant weight of Undaria sporophytes increased steadily over the experimental period, but it increased in four different ways. Undaria pinnatifida increased body weight by growth in length and width (October–early December), and by growth in width with the thickening of blade and stipe when sporophytes began to be fertile (December–January). In the middle of January, growth in length and width had almost stopped with the maturation of Undaria sporophytes. Finally, the weight of Undaria increased again by growth in width at the end of February. Present results indicate that Undaria sporophytes increase body weight by growth in length and width at different times, and the relationship between reproduction and vegetative growth is exclusive. Plant weight was positively correlated and fitted well with stipe width and blade width. The blade of Undaria was very thin (ca. 254 μm) and breakable by wave action, but its stipe was strong and relatively thick (ca. 8.7 mm). Furthermore, the fertility of U. pinnatifida was fitted better with stipe width than blade width. Thus, we suggest that the stipe width is the most feasible character with which to estimate the growth and reproduction of U. pinnatifida sporophytes in the cultivation farm.     

kLa of stirred tank bioreactors revisited

By means of improved feedback control k_La measurements become possible at a precision and reproducibility that now allow a closer look at the influences of power input and aeration rate on the oxygen mass transfer. These measurements are performed online during running fermentations without a notable impact on the biochemical conversion processes. A closer inspection of the mass transfer during cultivations showed that at least the number of impellers influences mass transfer and mixing: On the laboratory scale, two hollow blade impellers clearly showed a larger k_La than the usually employed three impeller versions when operated at the same agitation power and aeration rate. Hollow blade impellers are preferable under most operational conditions because of their perfect gas handling capacity. Mixing time studies showed that these two impeller systems are also preferable with respect to mixing. Furthermore the widths of the baffle bars depict a significant influence on the k_La. All this clearly supports the fact that it is not only the integral power density that finally determines k_La.     

Variations in the alginate content and composition of Durvillaea antarctica and D. willana from southern New Zealand

The brown seaweeds Durvillaea antarctica andD. willana are dominant components of the lowerlittoral and upper sublittoral of exposed rocky shoresin southern New Zealand. Tissue samples of bothspecies, harvested from a site on the south-east coastof South Island over a period of 2 years, wereanalysed for alginate content and composition.Individuals of both species were further separatedinto different blade (lamina and palm) and stipe(cortex and medulla) fractions to assess variationwithin the thallus. On average the alginate contentand frequency of mannuronic acid (F_m) was higherin D. antarctica than in D. willana. Blades contained more alginate than stipes, laminaeand stipes were rich in mannuronic acid whereasholdfasts were rich in guluronic acid. Variations incomposition are considered to reflect the functionaldifferences of the tissue, giving flexibility to bladeand stipe and rigidity to the holdfast. Despitefluctuations in content and composition betweencollection times no seasonal trends in eithercomponent were apparent.     

Ediacaran macroalgal holdfasts and their evolution: a case study from China

A macroalgal holdfast (root-like structure) anchored or grown into sediments is a key trait of metaphytes and eukaryotic algae. Various patterns and taphonomic variants of congeneric holdfasts are preserved on the bedding planes of black shales of the Ediacaran Wenghui biota in South China. The macroalgal holdfast, which commonly consists of a rhizome, rhizoid and pith (perhaps mechanical tissue), can be morphologically classified into ten types within four groups of rhizomes: bare rhizome holdfasts (Grypania, Tongrenphyton and Sectoralga-type holdfasts), canopy rhizome holdfasts (Gemmaphyton, Gesinella, Discusphyton and Baculiphyca-type holdfasts), pithy rhizome holdfasts (Zhongbaodaophyton and pithy-cone-type holdfasts) and differentiated rhizome holdfasts (Wenghuiphyton-type holdfasts). Analysis of the Precambrian macroalgal record indicates that rhizomes played a more important role in the evolution of macroalgal holdfast than rhizoids. The following evolutionary stages of development of macroalgal holdfasts are proposed: (1) growth of the basal thallus into sediments (Grypania-type holdfast); (2) development of a primitive (indistinct) rhizome from the base of a stipe or thallus (Tongrenphyton-type and Sectoralga-type holdfasts); (3) growth of a distinct rhizome with rhizoid (Gemmaphyton-type, Gesinella-type and Discusphyton-type holdfasts); (4) development of a pith in the stipe (Baculiphyca-type holdfast); (5) pith extension into the rhizome (Zhongbaodaophyton-type and pithy-cone-type holdfasts); and (6) rhizome differentiation and development of a complex holdfast system (Wenghuiphyton-type holdfast).     

The genus Lessonia Bory (Phaeophyta,Laminariales) in Southern Chile and Argentina

Study of Lessonia in southern Chile and Argentina indicates that it is represented there by three species: L. nigrescens Bory, L. flavicans Bory and L. vadosa Searles sp. nov. Plants of these three species have been confused with each other in the literature, but appear clearly separable. Lessonia nigrescens is distinguished from the others by its massive, solid holdfast and numerous stipes. In Lessonia flavicans and L. vadosa a single stipe arises from a branched holdfast. These two species are distinguishable from each other in details of blade morphology, anatomy, and ecology. The taxonomic status of plants in South America outside of the study area remains unresolved. Populations in the Falkland Islands and in northern Chile in particular are in need of further study.