Habitat-specific size structure variations in periwinkle populations (<Emphasis Type="Italic">Littorina littorea)</Emphasis> caused by biotic factors

Shell size distribution patterns of marine gastropod populations may vary considerably across different environments. We investigated the size and density structure of genetically continuous periwinkle populations (Littorina littorea) on an exposed rocky and a sheltered sedimentary environment on two nearby islands in the south-eastern North Sea (German Bight). On the sedimentary shore, periwinkle density (917 ± 722 individuals m⁻²) was about three times higher than on the rocky shore (296 ± 168 individuals m⁻²). Mean (9.8 ± 3.9 mm) and maximum (22 mm) shell size of L. littorea on the sedimentary shore were smaller than on the rocky shore (21.5 ± 4.2 and 32 mm, respectively), where only few small snails were found. Additionally, periwinkle shells were thicker and stronger on the rocky than on the sedimentary shore. To ascertain mechanisms responsible for differences in population structures, we examined periwinkles in both environments for growth rate, predation pressure, infection with a shell boring polychaete (Polydora ciliata) and parasitic infestation by trematodes. A crosswise transplantation experiment revealed better growth conditions on the sedimentary than on the rocky shore. However, crab abundance and prevalence of parasites and P. ciliata in adult snails were higher on the sedimentary shore. Previous investigations showed that crabs prefer large periwinkles infested with P. ciliata. Thus, we suggest that parasites and shell boring P. ciliata in conjunction with an increased crab predation pressure are responsible for low abundances of large periwinkles on the sedimentary shore while high wave exposure may explain low densities of juvenile L. littorea on the rocky shore. We conclude that biotic factors may strongly contribute to observed differences in size structure of the L. littorea populations studied on rocky and sedimentary shores.     

Are rocky shore ecosystems affected by nutrient-enriched seawater? Some preliminary results from a mesocosm experiment

The response of rocky shore ecosystems to increased nutrient availability was examined in eight land-based mesocosms designed for hard-bottom littoral communities built at Marine Research Station Solbergstrand (Norway). The average seawater volume in each basin was 9 m³ with an average water residence time of about 2 h. A tidal regime resembling that in the fjord was maintained in the basins, and waves were generated regularly. NH₄NO₃ and H₃PO₄, at a constant molar NP ratio of 16:1, was added into 6 basins at concentrations 1, 2, 4, 8, 16, 32 M DIN above the background DIN concentration during 1 1/2 years. Two mesocosms were kept as control treatment. Marine communities were introduced into the basins two years prior to the start of nutrient dosage. The effects of nutrient enrichment were few and only marginal during the first year of nutrient addition, while some effects became more obvious during the second year. The growth rate of the periphyton and fast-growing macroalgae communities was stimulated by nutrient enrichment, while the response was less evident among the perennial fucoids. The structure of the macroalgal communities, however, did not change during 16 months' measurements. In contrast, growth on artificial rock substrates during the same period of time revealed intensive growth of the fast-growing Ulva lactuca in high-dosed basins compared with low-dosed and control basins, which were dominated by the fucoid Fucus serratus. The fauna communities exhibited only a minor response to nutrient treatment. The common periwinkle Littorina littorea, however, appeared with increased abundance in the high-dosed basins. The total system metabolism tended to increase slightly, but not significantly, with increased nutrient loading.     

Human urine is an excellent liquid waste for the culture of fish food organism, Moina micrura

With a view to converting human urine into bio-wealth in the form of zooplankton, the nutrient potentials of liquid wastes (0.11 mL L⁻¹)—(i) human urine (♂), (ii) cow urine, (iii) human–cow mixed urine or some solid wastes (0.11 g L⁻¹): (iv) vermi-compost, (v) cow dung, (vi) poultry droppings and (vii) mixed wastes (vermi-cow-poultry)—were evaluated for the mass culture of zooplankton Moina micrura in 24 outdoor tanks (4500 L) in triplicate treatments using life table as indicator during the period of October–December, 2005. Neonates of Moina micrura held in the treatment with human urine started reproduction at least 4 days earlier than other solid wastes tested. Total number of Moina micrura enumerated in the culture tank, related with offspring production per life span, was maximum in case of human urine treatment, followed by human–cow mixed urine, cow urine, vermin-compost, poultry droppings, mixed wastes (vermin–cow–poultry), cow dung and control treatments. The relationship between the total offspring production per female per life span and the nitrogen content of water in different treatments implied that human urine was an excellent liquid waste that can be used for the mass production of zooplankton Moina micrura required for larval and post larval rearing of commercial fishes.     

Use of agricultural wastes for xanthan production by Xanthomonas campestris

Four different acid-hydrolyzed wastes, from melon, watermelon, cucumber and tomato were compared for xanthan production. Growth of Xanthomonas campestris, xanthan biosynthesis, kinetics and chemical composition were investigated. Both growth and xanthan production were dependent on the acid hydrolysate concentrations and available nitrogen. Melon acid hydrolyzed waste was the best substrate for xanthan production. Exopolysaccharide obtained throughout this study was compared to commercial xanthan, showing a very similar chemical composition. Acid hydrolyzed wastes are proposed as a new carbon source for xanthan production. Received 16 July 1998/ Accepted in revised form 8 October 1998     

Differentiation during asexual reproduction and regeneration in a microturbellarian

Affordable biological technology for the reclamation of wastes and water of the waste streams from intensive livestock units is important in a country short of water. This study tested the concept of reclamation of waste by Streptocephalus macrourus (Crustacea: Anostraca) from the effluent of a high rate algal pond processing livestock wastes. S. macrourus showed a growth efficiency of 39% to 74% when fed optimal rations and cultured at densities between 10 and 400 1^–1. The maximum daily growth rates (0.15–0.21) approximate the growth rates of cladoceran or rotifer cultures managed for maximal biomass production. S. macrourus' ability to withstand crowding enabled the production from S. macrourus cultures (up to 91.8 mg dry mass l^–1 d^–1, or 1241 mg wet mass l^–1 d^–1) to exceed production recorded from cladoceran or rotifer cultures. Temperature influenced growth rate, with the highest growth rate occurring at 24 °C. The dilution rate of continuously fed cultures influenced growth rate, with the optimum dilution rate tested being 10 ml organism ^–1 d^–1. Mass mortality occurred when organisms were held at a density of 4000 l^–1. S. macrourus is able to convert algae grown on livestock waste efficiently into anostracan biomass, and is able to give a very high daily production.     

Effects of water level fluctuation on the growth ofNelumbo nucifera Gaertn. in Lake Kasumigaura,Japan

Investigations were made of the growth ofNelumbo nucifera, an aquatic higher plant, in a natural stand in Lake Kasumigaura. A rise of 1.0 m in the water level after a typhoon in August 1986 caused a subsequent decrease in biomass ofN. nucifera from the maximum of 291 g d.w. m⁻² in July to a minimum of 75 g d.w. m⁻². The biomass recovered thereafter in shallower regions. The underground biomass in October tended to increase toward the shore. The total leaf area index (LAI) is the sum of LAI of floating leaves and emergent leaves. The maximum total LAI was 1.3 and 2.8 m² m⁻² in 1986 and 1987, respectively. LAI of floating leaves did not exceed 1 m² m⁻². The elongation rates of the petiole of floating and emergent leaves just after unrolling were 2.6 and 3.4 cm day⁻¹, respectively. The sudden rise in water level (25 cm day⁻¹) after the typhoon in August 1986 caused drowning and subsequent decomposition of the mature leaves. Only the young leaves were able to elongate, allowing their laminae to reach the water surface. The fluctuation in water level, characterized by the amplitude and duration of flooding and the time of flooding in the life cycle, is an important factor determining the growth and survival ofN. nucifera in Lake Kasumigaura.     

Pulp and paper mill solid wastes as substrates for single-cell protein production

We have investigated the upgrading of some typical pulp and paper mill solid wastes into protein-enriched animal feed using the cellulolytic fungus Chaetomium cellulolyticum. The waste residues used were six different primary clarifier sludges and a sample of tertiary centricleaner rejects. These were obtained from mills whose modes of operation spanned the range typically in present-day usage: groundwood, sulfite, semichemical, Kraft, and thermomechanical pulping, with and without bleaching. Crude protein production from the solid waste residues is compared to that obtainable from fermentation of untreated or caustic-pretreated sawdusts. Some of these waste residues, especially the Kraft pulp mill rejects, appear to be promising sources of substrate for single-cell protein production. In these preliminary findings, up to 28% dry weight crude protein content of the product has been obtained at specific growth rates of up to 0.12hr^?1 on direct utilization of the wastes.     

Improvement of the shallow water communities following reductions of industrial outlets and sewage discharge in the Hvaler estuary,Norway

Three years' investigations of the shallow water communities at 30 different rocky shore sites in an estuary in southeast Norway were carried out during 1980–1982. The flora was mostly poor, possibly due to a combination of fresh water exposure, industrial and sewage outlets. The aim of these investigations was to establish the environmental status of the estuary prior to huge outlet reductions in 1989/90. Later reinvestigations during 1992–1994, following a significant reduction of titanium dioxide wastes and building of a new chemical sewage treatment plant have shown an improvement of the macroalgal and benthic animal communities. In the formerly most polluted area, the number of benthic taxa has increased since 1980–1982 by 50–350% in 1992–1994.     

Continuous biomethanization of agrifood industry waste: A case study in Spain

The anaerobic digestion technology is a biological treatment widely used to reduce the pollution load of wet waste biomass. In this work we present the results obtained by performing extensive experiments of anaerobic digestion of slaughterhouse waste, tomato industry waste and olive oil industry waste in continuous mode, which were designed to demonstrate that anaerobic digestion is an effective technology from an environmental and economic point of view.Biogas yields obtained are between 35.22 and 5.45 Nm³ biogas/m³ olive oil industry waste and tomato industry waste respectively and the slaughterhouse wastes achieve intermediate production, 30.86 Nm³ biogas/m³ municipal slaughterhouse waste and 22.53 Nm³ biogas/m³ Iberian pig slaughterhouse waste. Moreover, it possible to degrade between 63.46 and 75.3% of the initial organic matter.If these results are analyzed, the environmental, energetic economic benefits of anaerobic digestion can be quantified. Biomethanation of all these wastes generated annually in Extremadura could prevent the emission of 134,772 t of equivalent carbon dioxide, generate an energy similar to that provided by 2826 toe and reach payback times from 3.29 to 3.75 years for anaerobic digestion plant designed to treat the wastes generated by a medium-sized industry. So, we have fulfilled all the planned aims.     

Distribution in relation to life history in the directdeveloping gastropod Batillaria cumingi (Batillariidae) on two shores of contrasting substrata

Population structure and spatial distribution with growth ofthe direct-developing gastropod, Batillaria cumingi, were investigatedon two shores of differing substrata. Sand-mud shore and rockyshore populations differed in size structure; first-year snailswere ca. 7 mm in shell length (SL) in both populations, whereassecond-year snails, merging with older cohorts, measured 15–25mm SL in the sand-mud shore population and ca. 15 mm SL in therocky shore population. Egg distribution matched adult distributionin the sand-mud shore population, but was more restricted thanthat of adults in the rocky shore population. The distributionof newly-hatched juveniles (0–1 mm SL) was restricted inboth populations, but the growth stage at which snails extendedtheir distribution differed between the two populations; 1–2.5mm SL on the sand-mud shore and 5 mm SL on the rocky shore. Floatingachieved by early juveniles (ca. 2 mm SL), was commonly observedin the sand-mud population, but rarely in the rocky shore population.The sudden expansion in distribution of the 1–2.5 mm SLgrowth stage in the sand-mud shore population is consideredto have been caused by floating, while expansion of the distributionof older growth stages (>5 mm SL) in the rocky shore populationprobably occurs by crawling. (Received 13 May 1998; accepted 25 August 1998)     

A yield control approach to assess phytoavailability of Zn and Cu in irradiated,composted sewage sludges and composted manure in field experiments: I. Zinc

The threat of spreading diseases is a serious concern when organic wastes are applied to farmland. Irradiation and composting are effective methods to reduce pathogens. Field experiments were conducted to assess the influence of these pathogen-eliminating methods on plant availability of Zn in the wastes. Four organic wastes: digested and dewatered (DSS), irradiated (DISS), composted (DICSS) sewage sludge and composted livestock manure (CLM) were applied during two growing seasons at 10, 20, 30, and 40 t solids ha^–1 year^–1. Available N and K in the wastes were estimated and N and K fertilizers were added to the soil to equalize available N and K supply among treatments to avoid dilution of crop Zn concentration. A control treatment (CT) received fertilizers but no waste. Lettuce, snap beans and petunias were grown in 1990, and two cuts of lettuce were harvested in 1991. The influence of waste Zn application on crop Zn concentration was studied within approximately equal crop yields. Crop Zn concentration increased in all crops treated with DSS or DISS, and often reached a maximum at the 30 t ha^–1 rate of application, then slightly decreased at 40 t ha^–1. The response of crop Zn concentration to the amount of Zn applied in the wastes was best described by quadratic equations. Waste application also significantly increased soil Zn availability index, which was a function of DTPA (diethylenetriamine pentaacetic acid)-extractable soil Zn and soil pH. The index was highly correlated with crop Zn concentration. Although Zn concentration in DICSS was similar to those in DSS and DISS, Zn applied in DICSS did not increase crop Zn concentration or soil availability index. Composting reduced the availability of Zn. The similar concentrations of Zn in DSS and DISS in both years allowed the use of a paired t-test to determine the differences in crop Zn concentration caused by application of DSS and DISS. Zinc applied in DISS produced a higher Zn concentration in bean pods than Zn applied in DSS (t > T _0.05 at P < 0.02, df = 15), indicating that irradiation increased phytoavailability of Zn in the sludge. However, no similar effect was found in Zn concentrations in the two cuts of lettuce in 1991 or in soil Zn availability index.     

Biofiltering efficiency, uptake and assimilation rates of Ulva clathrata (Roth) J. Agardh (Clorophyceae) cultivated in shrimp aquaculture waste water

The growth, biofiltering efficiency and uptake rates of Ulva clathrata were studied in a series of outdoor tanks, receiving waste water directly from a shrimp (Litopenaeus vannamei) aquaculture pond, under constant aeration and two different water regimes: (1) continuous flow, with 1 volume exchange a day (VE day^-1) and (2) static regime, with 1 VE after 4 days. Water temperature, salinity, pH, dissolved inorganic nitrogen (DIN), phosphate (PO₄), chlorophyll-a (chl-a), total suspended solids (TSS), macroalgal biomass (fresh weight) and tissue nutrient assimilation were monitored over 12 days. Ulva clathrata was highly efficient in removing the main inorganic nutrients from effluent water, stripping 70–82% of the total ammonium nitrogen (TAN) and 50% PO₄ within 15 h. Reductions in control tanks were much lower (Tukey HSD, P < 0.05). After 3 days, the mean uptake rates by the seaweed biomass under continuous flow were 3.09 mg DIN g DW day⁻¹ (383 mg DIN m⁻² day⁻¹) and 0.13 mg PO₄ g DW day⁻¹ (99 mg PO₄ m⁻² day⁻¹), being significantly higher than in the static regime (Tukey HSD, P < 0.05). The chl-a decreased in seaweed tanks, suggesting that U. clathrata inhibited phytoplankton growth. Correlations between the cumulative values of DIN removed from the water and total nitrogen assimilated into the seaweed biomass (r = 0.7 and 0.8, P < 0.05), suggest that nutrient removal by U. clathrata dominated over other processes such as phytoplankton and bacterial assimilation, ammonia volatilization and nutrient precipitation.     

Rotifer distributions in the coastal waters of the northeast Pacific Ocean

Absolute abundance of rotifers was assessed from 5 to 80 km across the continental shelf off of the southern Oregon coast (U.S.A.) in the northeast Pacific Ocean. A total of 97 vertically stratified water samples were collected at 49 stations from two depths, 3 and 30 m. Coastal upwelling conditions were indicated, with decreased temperature, increased salinity and higher chlorophyll-a concentrations closer to shore. Two rotifer genera, Synchaeta and Trichocercaoccurred within 16 km of shore with densities increasing closer to shore. Synchaeta reached densities of 64 inds l^–1 while Trichocerca was sparse (<1 inds. l^–1). Rotifers were most abundant at 3 m and the densest aggregations appear to be associated with estuary outlets, suggesting that estuaries may be important in exporting rotifers to nearshore coastal waters.     

Water relations and haemolymph composition of two intertidal spiders (order araneae)

Water relations and haemolymph composition have been compared in two intertidal spiders which inhabit different zones of a rocky shore in the Cape Peninsula. Evaporative water losses of the mid-shore species, Desis formidabilis (O.P. Cambridge), are much greater than those of Amaurobioides africanus Hewitt, which occurs higher on the shore. Both species avoid desiccating conditions by remaining in silk-lined nests in suitable microclimates, and the nests of Desis formidabilis are submerged for a substantial portion of each tidal cycle. Osmolarities and sodium, potassium, magnesium, and chloride concentrations were measured for the haemolymph of both spiders. The haemolymph chemistry of Amaurobioides africanus resembles that of terrestrial spiders. The haemolymph of Desis formidabilis, however, has an osmolarity of 930 mOsm/l, a sodium concentration of 451 mM/l and a chloride concentration of 466 mM/l. It is suggested that this unusually concentrated haemolymph may represent an adaptation to a diet of marine crustaceans.     

Production of butanol from starch-based waste packing peanuts and agricultural waste

We examined the fermentation of starch-based packing peanuts and agricultural wastes as a source of fermentable carbohydrates using Clostridium beijerinckii BA101. Using semidefined P2 medium containing packing peanuts and agricultural wastes, instead of glucose as a carbohydrate source, we measured characteristics of the fermentation including solvent production, productivity, and yield. With starch as substrate (control), the culture produced 24.7 g l⁻¹ acetone–butanol–ethanol (ABE), while with packing peanuts it produced 21.7 g l⁻¹ total ABE with a productivity of 0.20 g l⁻¹ h⁻¹ and a solvent (ABE) yield of 0.37. Cell growth in starch, packing peanuts, and agricultural wastes medium was different, possibly due to the different nature of these substrates. Using model agricultural waste, 20.3g l⁻¹ ABE was produced; when using actual waste, 14.8 g l⁻¹ ABE was produced. The use of inexpensive substrates will increase the economic viability of the conversion of biomass to butanol, and can provide new markets for these waste streams. Journal of Industrial Microbiology & Biotechnology (2002) 29, 117–123 doi: 10.1038/sj.jim.7000285 Received 14 November 2001/ Accepted in revised form 07 June 2002     

Distillery and curd whey wastes as viable alternative sources for biosurfactant production

Biosurfactant production from synthetic medium and industrial waste, viz. distillery and whey wastes was investigated by using an oily sludge isolate Pseudomonas aeruginosa strain BS2. In synthetic medium separately supplemented with glucose and hexadecane as water-soluble and -insoluble carbon sources, respectively, strain BS2 reduced the surface tension of the fermentation broth from 57 to 27 mN/m. The culture produced biosurfactant during the stationary growth phase and its yield was 0.97 g/l. The culture utilized distillery and whey wastes for its growth, as maximum cell counts reached to 54 × 10⁸ and 64 × 10⁹ c.f.u./ml from an initial inoculum size of 1 × 0⁵ c.f.u./ml, respectively, within 48 h of incubation and in these wastes the yields of biosurfactant obtained were 0.91 and 0.92 g/l, respectively. In synthetic medium, distillery and whey wastes, strain BS2 produced a crystalline biosurfactant which belonged to the category of secondary metabolites and its maximum production occurred after the onset of nitrogen-limiting conditions. After recovering biosurfactant from the fermented waste, the chemical oxygen demand (COD) of distillery and whey wastes was significantly reduced by 81 and 87%, respectively. Total acids, nitrogen and phosphate levels in distillery waste were reduced by 90, 92 and 92%, respectively, while in case of whey waste the concentration of these nutrients was reduced by 88, 95 and 93%, respectively. The isolated biosurfactant possessed potent surface active properties, as it effectively reduced the surface tension of water from 72 to 27 mN/m and formed 100% stable emulsions of a variety of water-insoluble compounds such as hydrocarbons, viz. hexadecane, crude oil, kerosene and oily sludge and pesticides, viz. dichlorodiphenyltrichloroethane (DDT) and benzene hexachloride (BHC). The effectiveness of biosurfactant was also evident from its low critical micellar concentration (CMC) which was 0.028 mg/ml.     

Nest site choice by the intertidal spider Desis formidabilis (Araneae: Desidae) and nest utilisation by its hymenopteran egg parasitoid

1. Echthrodesis lamorali Masner, 1968 is the only known parasitoid of the eggs of the intertidal rocky shore spider Desis formidabilis O.P. Cambridge 1890 and is endemic to a small area of South Africa. 2. The abundance of spider nests and parasitoid presence were assessed in relation to their in‐ and between‐shore location at multiple sites within the distribution of E. lamorali along the Cape Peninsula (Western Cape, South Africa). 3. Desis formidabilis nests were more abundant in the mid‐shore zone than higher up or lower down the shore. Spider population sizes also differed between collection sites, with higher numbers recorded on the cooler western coast of the peninsula. 4. Evidence of parasitoid activity was recorded in 43.31% of the 127 nests and 13.85% of the 592 egg sacs they contained. 5. Where parasitoids gained entry to a spider egg sac, oviposition took place into all of the eggs present. 6. Incidence of wasp activity was positively correlated with spider nest concentration, not with height up the shore, suggesting that both the host and parasitoid are tolerant of salt‐water inundation. 7. These results should assist managers of the Table Mountain National Park, in which the full distribution of E. lamorali falls, to better understand this component of rocky shore community dynamics.     

The vertical distribution of Chthamalus montagui and Chthamalus stellatus (Crustacea,Cirripedia) in two areas of the NW Mediterranean Sea

The Mediterranean Sea is characterised by a small tidal range (0.3–1 m). Despite this, intertidal communities are well established and their upper limits often extend above mean high water level. Organisms living in the intertidal region and in the supralittoral zone rely on both tides and wave action to perform their biological functions. Lack of food, desiccation and predation are common stresses in such a harsh environment. The present study deals with the vertical distribution of two species of intertidal barnacles, Chthamalus montagui Southward and Chthamalus stellatus (Poli), which are the main constituents of the barnacle belt along Mediterranean rocky shores. Previous work, carried out in the Atlantic, showed that where the distribution ranges of the two Chthamalus species overlap, C. montagui is more common in the upper barnacle zone while C. stellatus is dominant lower down. The main aims of our study are: (1) to establish if there is a relationship between position and extension of the barnacle belt on the shore and tidal range and/or wave exposure, (2) to test the hypothesis that in the study areas C. montagui is more abundant than C. stellatus high on the shore, and that the pattern is reversed lower down. Barnacle populations were monitored in summer 1998 in the Gulf of Genoa (Ligurian Sea) and in the Gulf of Trieste (North-Adriatic Sea). The two areas differ in tidal range and hydrodynamism, the former presenting quite strong wave action and a tidal range of 30 cm, the latter having limited wave action and 1 m tidal range. Three shores were randomly selected in each gulf and two transects on each shore. Counts of barnacles in 10 ^* 10 cm quadrats were done at different shore heights along each transect. The data was subjected to analysis of variance. Results showed that a more pronounced hydrodynamic regime corresponded to a shift of the barnacle belt towards the higher shore (Gulf of Genoa), while in more sheltered areas (Gulf of Trieste), the barnacle distribution was confined to the intertidal region. The relative spatial distribution of C. montagui and C. stellatus within the barnacle belt varied locally, even between transects on the same shore, and this obscured the distribution pattern along the vertical gradient. Nevertheless, it was still possible to conclude that at mid and high shore in Genoa, C. stellatus was more abundant than C. montagui, while in Trieste the pattern was reversed.     

EFFECTS OF SHADING BY ADULTS ON THE GROWTH OF BLADE-STAGE MACROCYSTIS PYRZFERA (PHAEOPHYTA) DURING AND AFTER THE 1982–1984 EL NIÑO1

The effect of shading by an adult canopy on blade-stage Macrocystis pyrifera (L.) C. A. Agardh was estimated by comparing the average growth rate of individuals under a canopy to that of individuals in a canopy gap. This comparison was made in 1983 during a strong El Niño and again in 1986 after the El Niño. Estimated nutrient concentrations in 1983 were two orders of magnitude below those in 1986, whereas ambient light levels were over 3 times higher. The kelp canopy caused similar proportional light reductions (20–30%) during both years. Blades grew 18% slower under the canopy than in the clearing in 1983 and about 77% slower under the canopy in 1986. Blade-stage individuals grew at the same rates in clearings in 1983 and 1986. Regardless of shading, the average growth rate of blade-stage kelp under the ambient, low-nutrient conditions of 1983 was higher than that later observed for multifronded juveniles during the same El Niño. The growth of blade-stage kelp was more like that of larger juveniles growing under high-nutrient conditions. The difference may be due to greater concentrations of nutrients very near the sea floor where single blades are growing compared to concentrations higher in the water column where larger kelp have most of their tissues.