Estimation of Primary Productivity by Chlorophyll a in vivo Fluorescence in Freshwater Phytoplankton

Primary productivity in marine waters is widely estimated by the measurements of ¹⁴C incorporation, the underwater light climate, and the absorption spectra of phytoplankton. In bio-optical models the quantum efficiency of carbon fixation derived from ¹⁴C incorporation rates, the photosynthetically absorbed radiation derived from the underwater light climate, and the phytoplankton absorption spectra are used to calculate time- and depth-integrated primary productivity. Due to the increased sensitivity of commercially available fluorometers, chlorophyll a in vivo fluorescence became a new tool to assess the photosynthetic activity of phytoplankton. Since fluorescence data yield only relative photosynthetic electron transport rates, a direct conversion into absolute carbon fixation rates is not possible. Here, we report a procedure how this problem can be adressed in freshwater phytoplankton. We adapted a marine bio-optical model to the freshwater situation and tested if this model yields realistic results when applied to a hypertrophic freshwater reservoir. Comparison of primary productivity derived from ¹⁴C incorporation to primary productivity derived from Chl a fluorescence showed that the conversion of fluorescence data into carbon fixation rates is still an unsolved problem. Absolute electron transport rates calculated from fluorescence data tend to overestimate primary production. We propose that the observed differences are caused mainly by neglecting the package effect of pigments in phytoplankton cells and by non-carbon related electron flow (e.g., nitrogen fixation). On the other hand, the ¹⁴C incorporation rates can be artificially influenced by "bottle effects", especially near the water surface, where photoinhibition, photorespiration, and Mehler reaction can play a major role.     

Growth model for raceway pond cultivation of Desmodesmus sp. MCC34 isolated from a local water body

Biofuel production by microalgae has the advantage of higher biomass productivity over land crops. The selection of potential microalgae depends on the growth in outdoor mass cultivation during different seasons, which can be predicted by a mathematical model. Here, freshwater green algae were isolated from a local water body in Pilani, Rajasthan, India (geographical coordinates: 28°22′N 75°36′E) and characterized by microscopy and ribosomal RNA analysis. The strain was submitted to the Indian Agricultural Research Institute's microbial culture collection (IARI, India) and identified as Desmodesmus sp. MCC34. This strain, along with a fresh water green algae (Chlorella minutissima), two marine green algae species (Dunaliella salina and Dunaliella tertiolecta) and two nitrogen fixing cyanobacteria (Nostoc muscorum and Anabaena doliolum), were screened for lipid productivity and growth kinetics under culture room and raceway pond conditions. Desmodesmus sp. MCC34 showed the highest specific growth rate (0.26 day^?1), biomass production (1.9 g L^?1) and lipid productivity (103 mg L^?1 day^?1). The optimal temperature and saturating light intensity for maximal growth of Desmodesmus sp. MCC34 were 35 °C and 75 μmol m^?2 s^?1 with molar extinction coefficient of 0.22 m² g^?1, respectively. Desmodesmus sp. MCC34 was then subjected to outdoor cultivation in a 20‐m long raceway pond for 18 days during March and November 2013. The areal biomass productivity and volumetric biomass productivity were 13946.23 kg ha^?1 year^?1 and 56.94 mg L^? 1day^?1 during the month of March, decreasing to 6262.28 kg ha^?1 year^?1 and 25.57 mg L^? 1day^?1 during the month of November. A mathematical model was constructed to explain the relationship between biomass production and growth parameters such as temperature, light intensity and nutrient concentration. The productivity values predicted with the proposed model correspond well with the experimental data, suggesting the validity of the model.     

Yearly variation in primary productivity of marine phytoplankton from Cabo Frio (RJ,Brazil) region

Light saturation curves of natural assemblages of phytoplankton at 4 stations in Cabo Frio coastal waters were examined and related to changes in environmental conditions, based on 263 experiments carried out weekly during two years. No differences in specific productivity at saturating light (Pm ^B ) between stations were detected. Global mean value of Pm ^B was 4.7 mgC mgChla ^–1 h^–1 with a range from 0.50 to 15.29. Significant seasonal variations were not observed. Correlation and regression analysis attributed most of Pm ^B variation to temperature, salinity, nutrients and the phaeopigment: chlorophyll-a ratio.     

Protozoan Colonization Rates and Trophic Status of Some Freshwater Wetland Lakes

Primary productivity, chlorophyll a, phosphorus and nitrogen nutrients, and other chemical and physical parameters were measured in 13 wetland lakes in northern lower Michigan. These lakes included several examples located in each of the four major wetland types—bogs, fens, marshes, and swamps. Of the four types, the brown-colored waters of the acid bog lakes generally had the highest levels of primary productivity, chlorophyll a, phosphorus, and nitrogen. Primary productivity correlated positively with water color, total-N, and NH₃-N (α≤ 0.05). By these measures, waters of the bogs were the most eutrophic of the four types of wetland lakes. These findings would seem to contradict the generally-held concept that “dystrophic” bog lakes are extremely oligotrophic. Protozoan colonization of artificial substrate islands was monitored at each wetland site. The correlation between protozoan colonization rates (G values in the MacArthur-Wilson noninteractive model) and primary productivity, measured by 24-h light and dark bottle incubations, was significant at the 95% confidence level (r= 0.850, P= 0.001) and with water color at the 90% confidence level (r= 0.599, P= 0.084). It was concluded that protozoan colonization rate was an excellent indicator of the trophic status of wetland lakes.     

Photosynthetic efficiency of Chlorella sorokiniana in a turbulently mixed short light‐path photobioreactor

To be able to study the effect of mixing as well as any other parameter on productivity of algal cultures, we designed a lab‐scale photobioreactor in which a short light path (SLP) of (12 mm) is combined with controlled mixing and aeration. Mixing is provided by rotating an inner tube in the cylindrical cultivation vessel creating Taylor vortex flow and as such mixing can be uncoupled from aeration. Gas exchange is monitored on‐line to gain insight in growth and productivity. The maximal productivity, hence photosynthetic efficiency, of Chlorella sorokiniana cultures at high light intensities (1,500 μmol m^?1 s^?1) was investigated in this Taylor vortex flow SLP photobioreactor. We performed duplicate batch experiments at three different mixing rates: 70, 110, and 140 rpm, all in the turbulent Taylor vortex flow regime. For the mixing rate of 140 rpm, we calculated a quantum requirement for oxygen evolution of 21.2 mol PAR photons per mol O₂ and a yield of biomass on light energy of 0.8 g biomass per mol PAR photons. The maximal photosynthetic efficiency was found at relatively low biomass densities (2.3 g L^?1) at which light was just attenuated before reaching the rear of the culture. When increasing the mixing rate twofold, we only found a small increase in productivity. On the basis of these results, we conclude that the maximal productivity and photosynthetic efficiency for C. sorokiniana can be found at that biomass concentration where no significant dark zone can develop and that the influence of mixing‐induced light/dark fluctuations is marginal. © 2010 American Institute of Chemical Engineers Biotechnol. Prog., 2010     

Chla-specific productivity of picophytoplankton not higher than that of larger phytoplankton off the South Shetland Islands in summer

Size-fractionated chlorophyll a (Chla)-specific productivity (μgC μgChla ⁻¹ h⁻¹) was measured at 11 stations off the northern coast of the South Shetland Islands during summer. The Chla-specific productivity of the 2- to 10 or 10- to 330-μm fraction was highest at 100% and 23% light depths. The Chla-specific productivity of the 2- to 10-μm fraction was generally highest, and that of the <2 or 10- to 330-μm fraction was sometimes highest at 12% and 1% light depths. Temperature was less than 3°C within the euphotic zone at all stations. The hypothesis of Shiomoto et al., according to which Chla-specific productivity of picophytoplankton (<2 μm) is not significantly higher than that of larger phytoplankton (>2 μm) in water colder than 10°C, was supported on condition that light is not limited for larger phytoplankton. Received: 16 September 1997 / Accepted: 8 December 1997     

Mineralization of N and P along a trophic gradient in a freshwater mire

Release of inorganic nitrogen and phosphorus in the soil of a peatland (fen) in The Netherlands was measured by means of an in situ incubation technique. Three sampling stations were chosen along a gradient in the plant productivity and water chemistry of the fen. The station with the highest biomass production was located near the ditch that supplied the fen with water in amounts matching water losses through evaporation and downward percolation to the groundwater. Water chemistry at this station strongly resembled that of the ditch water. The two stations remote from the ditch had much lower plant biomass, and significantly lower pH, conductivity, and calcium and bicarbonate concentrations. The vegetation at these two stations was characterized by a thick Sphagnum carpet.The release of inorganic N and P was much faster at the two stations remote from the ditch than at that located near the ditch. The differences in mineralization rate are probably due to the differences in water chemistry; phosphates are more soluble at low than at high pH. The fast N mineralization at stations with a thick Sphagnum carpet may be related to the chemical composition of Sphagnum litter. The difference in productivity is not explained by the N and P mineralization rates. Direct supply of N and P from the ditch are probably the main cause of the high productivity at the station bordering the ditch.     

Zooplankton species diversity in Lake St. Clair,Ontario, Canada

Zooplankton species diversity and selected chemical parameters were investigated at three stations in Lake St. Clair, Ontario, Canada, from 15 June–26 August, 1971. Primary productivity and zooplankton species diversity were greatest at stations 1 and 2 which were enriched by the Thames River drainage. No significant correlation between total zooplankton diversity and chlorophyll a was found, however, within the Cladocera and Copepoda, positive correlations with chlorophyll a, reactive silicate and nitrate were shown at stations 2 and 3. Rotifer species diversity showed negative correlation with chlorophyll a, nitrate and reactive silicate.     

Relationship between small-scale structural variability and simulated vegetation productivity across a regional moisture gradient in southern Africa

The observed variability in vegetation structure within landscapes was used as the basis for model estimates of the range of annual productivity of landscape patches at four sites along a moisture gradient in southern Africa ranging from 879 to 365 mm mean annual rainfall. Principal components of patch‐scale variability in leaf area, woody biomass and vertical leaf profiles were derived from intensive characterization of the small‐scale spatial structure of woody vegetation at each site. For each site, the mean and extremes of the principal component distribution parameterized an ecophysiology model of vegetation productivity. Vegetation was most heterogeneous at intermediate locations along the rainfall gradient. Variability in vegetation structure led to a range of annual productivity within one site (600 mm) that accounted for 68% of the total range in mean productivity across all sites. Patch‐scale estimates of tree productivity were found to be primarily correlated to annual rainfall (r²=0.66, P=0.001) and not woody leaf area (r²=0.01, P=0.75), while grass productivity was found to be related to values of woody leaf area (r²=0.77, P<0.001) and not annual rainfall (r²=0.11, P=0.29). This result indicates that life‐form interactions have a significant role in controlling vegetation productivity across the rainfall gradient. The findings of this study emphasize the importance of considering heterogeneity rather than mean structure when modeling productivity, particularly when considering dynamic vegetation structure, where differences between landscape patches may not be well represented in the mean structure.     

Do light/dark cycles of medium frequency enhance phytoplankton productivity?

It has been suggested that turbulence with the resultant light/dark cycle and light gradient through which phytoplankton move, enhances their productivity. The stationary bottle incubation technique for estimating rates of primary productivity has mainly been criticized because of bottle effects, the elimination of natural turbulence and the presence of photo-inhibition. In a series of experiments where productivity was measured over static profiles and compared to the productivity in a mixed system, no definite conclusion could be reached regarding the effect of varying light/dark cycles of medium frequency (seconds to minutes). It appeared as though the ratio of the euphotic depth to mixing depth (Z _eu/Z _m) influenced productivity more than the duration of the light/dark cycle. The static bottle incubation method gave higher integral productivities than the mixed samples at low ratio's ofZ _eu/Z _m. It is suggested that mixing has two separate, but synergistic effects i.e. it not only moves the phytoplankton cells through a light/dark cycle, but also decreases the boundary layer, which increases the rate of exchange through the cell wall of nutrients and metabolites. In doing so more nutrients are available and light could be utilized more efficiently and therefore, productivity is increased.     

A preliminary analysis of the correlation of food-web characteristics with hydrology and nutrient gradients in the southern Everglades

We estimated trophic position and carbon source for three consumers (Florida gar, Lepisosteus platyrhincus; eastern mosquitofish, Gambusia holbrooki; and riverine grass shrimp, Palaemonetes paludosus) from 20 sites representing gradients of productivity and hydrological disturbance in the southern Florida Everglades, U.S.A. We characterized gross primary productivity at each site using light/dark bottle incubation and stem density of emergent vascular plants. We also documented nutrient availability as total phosphorus (TP) in floc and periphyton, and the density of small fishes. Hydrological disturbance was characterized as the time since a site was last dried and the average number of days per year the sites were inundated for the previous 10 years. Food-web attributes were estimated in both the wet and dry seasons by analysis of δ¹⁵N (trophic position) and δ¹³C (food-web carbon source) from 702 samples of aquatic consumers. An index of carbon source was derived from a two-member mixing model with Seminole ramshorn snails (Planorbella duryi) as a basal grazing consumer and scuds (amphipods Hyallela azteca) as a basal detritivore. Snails yielded carbon isotopic values similar to green algae and diatoms, while carbon values of scuds were similar to bulk periphyton and floc; carbon isotopic values of cyanobacteria were enriched in C₁₃ compared to all consumers examined. A carbon source similar to scuds dominated at all but one study site, and though the relative contribution of scud-like and snail-like carbon sources was variable, there was no evidence that these contributions were a function of abiotic factors or season. Gar consistently displayed the highest estimated trophic position of the consumers studied, with mosquitofish feeding at a slightly lower level, and grass shrimp feeding at the lowest level. Trophic position was not correlated with any nutrient or productivity parameter, but did increase for grass shrimp and mosquitofish as the time following droughts increased. Trophic position of Florida gar was positively correlated with emergent plant stem density.     

SEASONAL PRODUCTIVITY OF TWO RED ALGAE IN A CENTRAL CALIFORNIA KELP FOREST1

The in situ productivity of Botryocladia pseudodichotoma (Farl.) Kyl. and Rhodymenia californica var. californica Kyl., two common understory macrophytes in Macrocystis pyrifera (L.) C. Ag. forests, was determined once per month for one year. Gross productivity for B. pseudodichotoma was highest in October (1.23 mgC · g dry wt.^?1· h^?1), but rates were generally greater in spring and lower in summer. Respiration was variable throughout the year, especially in proportion to gross productivity. Gross productivity for R. californica var. californica was also highest in October (4.62 mgC · g dry wt.^?1· h^?1), and lowest in summer. Respiration was highly variable throughout the year. Deep (11 m) populations o/B. pseudodichotoma had 50% higher productivity than shallow (3 m) populations when incubated in shallow water, and 45% higher productivity when incubated in deep water. Populations of B. pseudodichotoma growing in shade at 3 m had 77% higher productivity than populations growing in sun at the same depth when incubated in a sun exposed location at 3 m. Respiration of the shade-adapted plants was only one-half that of the sun-adapted plants. In comparison with similar studies in other sub-tidal communities, net productivity of these two understory red algae is somewhat lower. Lack of strong seasonal productivity patterns and the highly variable underwater light regime suggests that understory algae may be adapted for rapid growth during short periods of high light, regardless of the time of year.     

Modeling algal productivity in large outdoor cultures and waste treatment systems

A deterministic mathematical model was used to describe the production of green microalgae (Scenedesmus obliquus and Coelastrum sphaericum) in outdoor mass cultures. The model was calibrated against 16 months of temperature and irradiance measurements, during which time productivity measurements were made in up to five ponds with surface areas of up to 263 m². During this period rates of algal dry matter production varied between 1·7 and 16·92 g m⁻² day⁻¹. The model predicted productivity to within 4·2% of the observed rates, for the same period. Negative productivity values (loss of biomass) were calculated for the months from November to January. It was concluded that appreciable amounts of biomass could be produced for 7 months per year in temperate areas.Several assumptions were made during the construction of the model, especially with regard to loss factors, such as: respiration, release of exuded organic carbon and photo-inhibition. The latter was included as a separate factor in the model and is merely conceptual. Several applications of the model are discussed, one of which concerns the relation between areal density and productivity, where the optimal areal density for maximal productivity was calculated to be 38–41 g (dry wt) m⁻². A distinction was also made between cultures which were mainly autotrophic and waste systems. It was shown that the presence of gilvin and/or tripton would adversely influence productivities and that the contribution of these factors to vertical light attenuation would have to be measured in waste systems.     

遥感GPP模型在中亚干旱区4个典型生态系统的适用性评价

总初级生产力(GPP)是全球生态系统碳循环的重要组成部分,对全球气候变化有重要影响。目前有多种遥感模型可以模拟总初级生产力,比较不同遥感模型在中亚干旱区上的适用性对推进全球干旱区碳收支估算具有重要意义。基于涡度协相关技术观测的四个地面站数据验证MOD17、VODCA2、VPM、TG、SANIRv五种模型的模拟精度。结果表明：(1)基于光能利用率理论的MOD17、VPM模型模拟咸海荒漠植被和阜康荒漠植被GPP的精度最高(R²分别为0.52和0.80),但在模拟草地、农田生态系统生产力时存在较明显的低估(RE>20%);基于植被指数的遥感模型TG模型、SANIRv模型模拟巴尔喀什湖草地生态系统和乌兰乌苏农田生态系统GPP的精度最高(R²分别为0.91和0.81),同时模拟值与实测值的相对误差也较低;基于微波的VODCA2模型模拟各生态系统生产力的效果最差。(2)水分亏缺是限制植被GPP的主要因素,因此是否合理考虑水分胁迫是影响GPP模型在中亚干旱区适用性的重要因素。研究揭示了遥感GPP模型在中亚干旱区的应用潜力,为推进全球植被碳通量的准确估...     

Sustainable, High-Yielding Outdoor Mass Cultures of Chaetoceros muelleri var. subsalsum and Isochrysis galbana in Vertical Plate Reactors

Continuous cultures of Chaetoceros muelleri and Isochrysis galbana were grown outdoors in flat plate-glass reactors in which light-path length (LPL) varied from 5 to 30 cm. High daily productivity (13 to 16 g cell mass per square meter of irradiated reactor surface) for long periods of time was obtained in reactors in which the optical path as well as cell density were optimized. 'Twenty centimeters was the optimal LPL, yielding the highest areal productivity of cell mass (g m^–2d^–1), eicosapentaenoic acid, and docosahexaenoic acid, which was identical with that previously found for polysaccharide production of Porphyridium and not far from the optimal LPL affecting maximal productivity in Nannochloropsis species. Relating the energy impinging on a given reactor surface area to the appropriate number of cells showed that the most efficient light dose per cell, obtained with the 20-cm LPL reactor, was approximately 2.5 times lower than the light dose available per cell in the 5-cm LPL reactor, in which a significant decline in areal cell density accompanied the lowest areal output of cell mass. The most effective harvesting regimen was in the range of 10% to 15% of culture volume harvested daily and replaced with fresh growth medium, resulting in a sustainable culture density of 24 × 10⁶ and 28 × 10⁶ cells/ml of C. muelleri and I. galbana, respectively.     

The cryptoendolithic microbial environment in the Ross Desert of Antarctica: Light in the photosynthetically active region

The vertical zonation of the Antarctic cryptoendolithic community appears to form in response to the light regime in the habitat. However, because of the structure of the habitat, the light regime is difficult to study directly. Therefore, a mathematical model of the light regime was constructed, which was used to estimate the total photon flux in different zones of the community. Maximum fluxes range from about 150m photons m^–2 s^–1 at the upper boundary of the community to about 0.1m photons m^–2 s^–1. Estimates of the annual productivity in the community indicate that the lowest zone of the community is light limited, with the maximal annual carbon uptake equivalent to less than the carbon content of one algal (Hemichloris) cell.     

Changes in specific photosynthetic rate of oceanic phytoplankton from the northeast Pacific Ocean

The study is based on data (n=244) from light-saturation experiments utilizing artificial incubation under fluorescent light. Values of maximum photosynthetic rate,P _max, and the light intensity at which it takes place,I _max, are estimated by non-linear regression using stepwise Gauss-Newton iterations. Estimated values ofP _max ranged from 0.85 to 5.48 mg C (mg Chla·h)^?1;I _max varied from 2.35 to 5.52 cal (cm²·h)^?1. The effects of time (months) and depth (illumination levels) and their interaction are evaluated by analysis of covariance using a linear model. A significant time-depth interaction is noted: The maximum specific primary productivity occurred in the surface layers during March, at the 50% light level during April, and at 1% level during May. Estimates ofP _max from simulated in situ primary productivity experiments for the same period are lower than those from light-saturation experiments. A comparison of data from light-saturation and simulated in situ experiments indicated that effects of duration of experiments and the quality of available light may affect primary productivity data considerably.     

Production in the Sea of Okhotsk

Primary production, microbial production and the density of planktonic microheterotrophs were estimated at 40 stations in the Okhotsk Sea in July-August 1992 during the seasonal phytoplankton minimum. The primary production by phytoplankton remained rather high even during this minimum. At most stations it was >0.6-0.8 g m^-2 day^-1, and in leftover patches of spring diatom 'bloom' it reached >5 g C m^-2 day^-1. The deep maxima of phytoplankton at the upper boundary of the seasonal thermocline were an ordinary phenomenon. The depth of the euphotic zone was normally 30-50 m in the open sea and 12-25 m at the shelf station. Any correlations between the phosphate contents in the upper mixed layer and primary production were absent at the stations. There was no adaptation of the phytoplankton to the light deficiency in deep maxima layers. The total numbers of bacterioplankton were 1-1.5 x 10⁶ ml^-1 and its biomass was close to 100 mg m^-3 in the open sea. All these numbers were 2-3 times greater at the shelf stations. In deep waters, the bacterioplankton biomass decreased to 10-40 mg m^-3. The microbial production in the upper layer was high, at 50-100 mg m^-3, decreasing 50-100 times in the deep waters. The numbers of ciliates in the upper water layer varied from 3 to 6 x 10³ l^-1 and were 1.5-2 times greater than in the shelf areas. Ciliate biomass was 60-100 mg m^-3 in the upper mixed layer, and per square metre varied to 1.5-2.5 g. The dominant ciliate taxa belonged to the naked oligotrichid genera Strombidium and Tontonia. Tentative calculations were made of the basin's annual primary production and for the analysis of energy balance in the ecosystem.      

Production by Hexagenia limbata in a warm-water reservoir and its association with chlorophyll content of the water column

We investigated the productivity of nymphs of the mayfly Hexagenia limbata in Lake Waco, a central Texas reservoir, and assessed its association with chlorophyll content of the water. We hypothesized that food availability measured as chlorophyll content of the water may directly associate with growth of Hexagenia and predict population productivity. To test this, we compared production by mayfly populations at two stations in the same reservoir; a northern station receiving water input with high chlorophyll content, and a southern station receiving water with low chlorophyll content. Both stations had similar substrate type and abundant mayflies. Benthic samples were collected from October 1984 through September 1985, and dissolved oxygen and temperature of the water were monitored.Annual production (size-frequency method) was 1270 mg m^–2 (P/B = 7.5) at the northern station and 1990 mg m^–2 (P/B = 6.1) at the southern station. The mean standing crop was 323 mg m ^{– 2} at the southern station and 169mg m^–2 at the southern station. Densities of mayflies at the two stations were not significantly different.Mean chlorophyll concentration (total mg pigment) during the sampling period was 23.5 mg m^–3 at the northern station and 16.7 mg m ^{– 3} at the southern station. Therefore, the station with lower mean chlorophyll content had higher secondary productivity by Hexagenia. Conversely, the station with higher mean chlorophyll content had lower mayfly productivity. The productivity of the mayfly populations did not positively associate with the chlorophyll content of the water, and chlorophyll content did not predict the success of the population of Hexagenia. Variation in mayfly growth success was associated with differences in temperature and dissolved oxygen. The northern station with higher chlorophyll content and lower productivity had low dissolved oxygen and temperatures higher than optimum for growth.     

Autoradiographic localization of 3H-γ-aminobutyric acid uptake in the lamina ganglionaris of Musca and Drosophila

Summary The uptake of ³H-GABA in the visual system of half-head preparations of Musca and Drosophila was studied by means of light and electron microscope autoradiography. Of all three ganglia, only the first synaptic region, the lamina ganglionaris, showed accumulation of radioactive grains, and there a preferential glial uptake could be found. Under normal light conditions at incubation (constant light flux of 100 Lux) the maximum of radio-activity was found in the marginal glia cells. Increasing the time of incubation produced also an increase in the number of grains per surface unit in the marginal glia cells. After changing the light intensity during incubation, quantitative modifications of the distribution of radio-activity were observed: incubating with stroboscopic illumination, the number of grains diminished in the marginal glia cells and remained constant in the epithelial cells; incubated in darkness, the epithelial cells became more intensely labelled whilst the number of grains decreased in the marginal cells.The possibility is discussed that the receptor axons 1–6 are the neurological elements of the lamina which use GABA as a transmitter. This hypothesis is lent some support from results of similar experiments with neurological mutants of Drosophila. In opm 18 there was a delayed uptake of ³H-GABA whereas in opm 3 and ebony the results were comparable to those found in Musca incubated in darkness. Behavioral studies on these mutants have demonstrated a defect, most probably related to the receptor system 1–6.