Housing areas — factors that need to be considered in the site layout and design process

The lack of any particular character in much new low‐rise housing is seen as resulting in the lack of attention paid to the detailing of space between buildings. The landscape architect's role in overcoming “placelessness” is considered, and the information needs of designers is discussed. The paper also offers some notes for the designer on user needs.     

Adaptation strategies of the corallimorpharian <Emphasis Type="Italic">Rhodactis rhodostoma</Emphasis> to irradiance and temperature

Corallimorpharians may dominate some habitats on coral reefs and compete with stony corals for access to light, yet little is known concerning their photosynthetic traits. At Eilat in the northern Red Sea, we observed that the abundance of individuals of the corallimorpharian Rhodactis rhodostoma decreased significantly with depth on the reef slope. Field and laboratory experiments revealed that they employ several mechanisms of photoadaptation to high irradiance on the shallow reef flat. Their endosymbiotic microalgae (zooxanthellae) varied significantly in both abundance and chlorophyll content with level of irradiance. Use of a diving pulse amplitude modulated fluorometer revealed that the zooxanthellae of R. rhodostoma effectively disperse excess light energy by expressing significantly higher values of non-photochemical quenching and maximum excitation pressure on photosystem II when experimentally exposed to high light (HL) versus low light (LL). Host corallimorpharian tissues mediated this response by shielding the algal symbionts from high irradiance. The endoderm of host tentacles thickened significantly and microalgal cells were located further from the mesoglea in HL than in LL. The clades of zooxanthellae hosted by the corallimorpharians also varied with depth. In shallow water, all sampled individuals hosted clade C zooxanthellae, while in deep water the majority hosted clade D. The photosynthetic output of individuals of R. rhodostoma was less affected by HL than was that of a stony coral examined. When exposed to both high temperature (HT) and HL, individuals of R. rhodostoma reduced their maximum quantum yield, but not when exposed to HL at low temperature (LT). In contrast, colonies of the scleractinian coral Favia favus reduced their photosynthetic output when exposed to HL in both temperature regimes. After 2 weeks of HT stress, R. rhodostoma polyps appeared to bleach completely but re-established their zooxanthella populations upon return to ambient temperature. We conclude that mechanisms of photoadaptation to high irradiance employed by both the endosymbiotic zooxanthellae and host corallimorpharians may explain in part the abundance of R. rhodostoma on some shallow reef flats. The ability to survive for weeks at HT while bleached also may allow corallimorpharians to repopulate shallow reef areas where scleractinians have been killed by thermal stress. B. Kuguru and G. Winters contributed equally to this work.     

Photosynthesis in air and in water of Acanthophora najadiformis growing within a narrow zone of the intertidal

In this work, we investigated photosynthetic gas exchange responses of the red alga Acanthophora najadiformis (collected in 1991 from a shore near Netanya, Israel) to heating, light, inorganic carbon and desiccation of the thalli. This was done to elucidate some of the contributing causes for its particular restricted vertical distribution within the lower intertidal zone of the Israeli Mediterranean rocky shore. In contrast to other common intertidal species, the photosynthetic performance of A. najadiformis was very sensitive to desiccation and to high temperature, and these factors may thus prevent the alga from growing above the surge zone. On the other hand, this species photosynthesized five times faster in air than in water (provided it was kept highly hydrated), and this may be the reason why it does not succeed in competing with other algae in the subtidal which use HCO ₃ ^- more efficiently.This paper is in partial fulfillment of a Ph.D. study by R. Einav in cooperation with the Department of Ecology, University of Bielefeld (under supervision of Prof. Dr. S. W. Breckle), under a cooperative arrangement between Tel Aviv and Bielefeld Universities     

Photosynthetic utilisation of inorganic carbon by seagrasses from Zanzibar, East Africa

Photosynthetic rates of eight seagrass species from Zanzibar were limited by the inorganic carbon composition of natural seawater (2.1 mM, mostly in the form of HCO₃ ⁻), and they exhibited more than three time higher rates at inorganic carbon saturation (>6 mM). The intertidal species that grew most shallowly, Halophila ovalis, Halodule wrightii and Cymodocea rotundata, showed the highest affinity for inorganic carbon (K _1/2 = ca. 2.5 mM), followed by the subtidal species (K _1/2 > 5 mM). Photosynthesis of H. wrightii, C. rotundata, Cymodocea serrulata and Enhalus acoroides was >50% inhibited by acetazolamide, a membrane-impermeable inhibitor of carbonic anhydrase, indicating that extracellular HCO₃ ⁻ dehydration is an important part of their inorganic carbon uptake. Photosynthetic rates of H. wrightii, Thalassia hemprichii, Thalassodendron ciliatum, C. serrulata and E. acoroides were strongly reduced by changing the seawater pH from 8.2 to 8.6 in a closed system. In H. ovalis, C. rotundata and Syringodiumisoetifolium, photosynthesis at pH 8.6 was maintained at a higher level than could be caused by the ca. 30% CO₂ concentration which remained in the closed experimental systems at that pH, pointing toward HCO₃ ⁻ uptake in those species. It is suggested that the ability of H. ovalis and C. rotundata to grow in the high, frequently air-exposed, intertidal zone may be related to a capability to take up HCO₃ ⁻ directly, since this is a more efficient way of HCO₃ ⁻ utilisation than extracellular HCO₃ ⁻ dehydration under such conditions. The inability of all species to attain maximal photosynthetic rates under natural conditions of inorganic carbon supports the notion that seagrasses may respond favourably to any future increases in marine CO₂ levels. Received: 19 March 1997 / Accepted: 31 March 1997     

Photosynthetic carbon acquisition in the red alga Gracilaria conferta

In this continuing study on photosynthesis of the marine red alga Gracilaria conferta, it was found that ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) in crude extracts had a K _m (CO₂) of 85 M. Since seawater contains only ca. 10 M CO₂, it appears that this alga must possess a CO₂ concetrating system in order to supply sufficient CO₂ to the vicinity of the enzyme. Because this species is a C₃ plant (and thus lacks the C₄ system for concentrating CO₂), but can utilize HCO₃ ^- as an exogenous carbon source, we examined whether HCO₃ ^- uptake could be the initial step of such a CO₂ concetrating system. The surface pH of G. conferta thalli was 9.4 during photosynthesis. At this pH, estimated maximal uncatalyzed HCO₃ ^- dehydration (CO₂ formation) within the unstirred layer was too slow to account for measured phostosynthetic rates, even in the presence of an external carbonic anhydrase inhibitor. This observation, and the marked pH increase in the unstirred layer following the onset of light, suggests that a HCO₃ ^- transport system (probably coupled to transmembrane H⁺/OH^- fluxes) operates at the plasmalemma level. The involvement of surface-bound carbonic anhydrase in such a system remains, however, obscure. The apparent need of marine macroalgae such as G. conferta for CO₂ concentrating mechanisms is discussed with regard to their low affinity of Rubisco to CO₂ and the low rate of CO₂ supply in water. The close similarity between rates of Rubisco carboxylation and measured photosynthesis further suggests that the carboxylase activity, rather than inorganic carbon transport and intercoversion events, could be an internal limiting factor for photosynthetic rates of G. conferta.     

Oxygen dynamics and transport in the Mediterranean sponge Aplysina aerophoba

The Mediterranean sponge Aplysina aerophoba kept in aquaria or cultivation tanks can stop pumping for several hours or even days. To investigate changes in the chemical microenvironments, we measured oxygen profiles over the surface and into the tissue of pumping and non-pumping A. aerophoba specimens with Clark-type oxygen microelectrodes (tip diameters 18–30 μm). Total oxygen consumption rates of whole sponges were measured in closed chambers. These rates were used to back-calculate the oxygen distribution in a finite-element model. Combining direct measurements with calculations of diffusive flux and modeling revealed that the tissue of non-pumping sponges turns anoxic within 15 min, with the exception of a 1 mm surface layer where oxygen intrudes due to molecular diffusion over the sponge surface. Molecular diffusion is the only transport mechanism for oxygen into non-pumping sponges, which allows total oxygen consumption rates of 6–12 μmol cm⁻³ sponge day⁻¹. Sponges of different sizes had similar diffusional uptake rates, which is explained by their similar surface/volume ratios. In pumping sponges, oxygen consumption rates were between 22 and 37 μmol cm⁻³ sponge day⁻¹, and the entire tissue was oxygenated. Combining different approaches of direct oxygen measurement in living sponges with a dynamic model, we can show that tissue anoxia is a direct function of the pumping behavior. The sponge-microbe system of A. aerophoba thus has the possibility to switch actively between aerobic and anaerobic metabolism by stopping the water flow for more than 15 min. These periods of anoxia will greatly influence physiological variety and activity of the sponge microbes. Detailed knowledge about the varying chemical microenvironments in sponges will help to develop protocols to cultivate sponge-associated microbial lineages and improve our understanding of the sponge-microbe-system.     

Photosynthetic utilisation of carbon and light by two tropical seagrass species as measured in situ

In situ measurements of seagrass photosynthesis in relation to inorganic carbon (Ci) availability, increased pH and an inhibitor of extracellular carbonic anhydrase were made using an underwater pulse amplitude modulated (PAM) fluorometer. By combining the instrument with a specially designed Perspex chamber, we were able to alter the water surrounding a leaf without removing it from the growing plant. Responses to Ci within the chamber showed that subtidal plants of the seagrasses Cymodocea serrulata and Halophila ovalis had photosynthetic rates that were limited by the ambient Ci concentration depending on the irradiance that was available during short-term photosynthesis–irradiance trials. Relative electron transport rates (RETRs) at light saturation (up to 500 μ mol photons m⁻² s⁻¹) increased by 66–100% when the Ci concentration was increased from ca. 2.2 to 6.2 mM. On the other hand, intertidal plants of the same species exhibited a much lesser limitation of photosynthesis by Ci at any irradiance (up to 1500 μ mol photons m⁻² s⁻¹). Both species were able to use HCO⁻ ₃ efficiently, and there was stronger evidence for direct uptake of HCO⁻ ₃ rather than extracellular dehydration of HCO⁻ ₃ to CO₂ prior to Ci uptake. Subtidally, H. ovalis and C. serrulata grew to 10 and 12 m, respectively, where ambient irradiances were approximately 16 and 11% of those at the surface. Maximum RETRs (at light saturation) were lower for these deep-growing plants than for the intertidally growing ones. For both species, the onset of light saturation of photosynthesis (E _k) occurred at approximately 100 μ mol photons m⁻² s⁻¹ for the deep water populations, which was four and two times lower than for the shallow populations of C. serrulata and H. ovalis, respectively. This, and the differences in maximal photosynthetic rates (RETR _max), reflects an acclimation of the deep-growing populations to the lower light environment. The results presented here show that photosynthesis, as measured in situ, was limited by the availability of Ci for the deeper growing plants in Zanzibar, while the intertidally growing plants photosynthesised at close to Ci saturation. The latter result is contrary to previous conclusions regarding Ci limitations for these intertidal plants, and, in general, our findings highlight the need for performing similar experiments in situ rather than under laboratory conditions. Received: 4 April 2000 / Accepted: 31 August 2000     

Use of pulse amplitude modulated (PAM) fluorometry for in situ measurements of photosynthesis in two Red Sea faviid corals

Measurements of the photosynthetic activity of symbiotic zooxanthellae in corals under natural growth conditions has been limited until recently, and this is one of the first reports on utilising a newly developed underwater pulse amplitude modulated (PAM) fluorometer (the Diving-PAM, Walz Gmbh, Germany) for such studies in situ. Photosynthetic responses to irradiance (photosynthetic photon flux, PPF) of the two faviid corals Favia favus (Forskål) and Platygyra lamellina (Ehrenberg) were measured while snorkelling or SCUBA diving (in August 1997), and we report here the results in terms of effective quantum yields of photosystem II (Y?) and estimated rates of photosynthetic electron transport (ETR, calculated as Y?×?0.5?× PPF?×?FA, where FA is the estimated fraction of light absorbed by the photosymbiont-containing tissue). Both species showed a reduction in Y with increasing actinic irradiances produced by the instrument above 500?μmol photons m^?2 s^?1, and the corresponding ETR values yielded apparently typical photosynthesis versus irradiance (P-I?) curves, which saturated between 1500 and 2000?μmol photons m^?2 s^?1. It was found that 30?s irradiation at each PPF level was sufficient to give optimal ETR values and, therefore, each P-I curve could be obtained within a few minutes. In situ point measurements from various areas of colonies under ambient light showed average ETR values within the range expected from the P-I curves. In order to test the Diving-PAM in an eco-physiologically relevant experiment, photosynthetic ETR versus PPF was measured for three sections of a large P. lamellina, each section of which received different natural irradiance levels. The results clearly demonstrated adaptations to the ambient light field in that vertical and downward-facing portions of the colony showed gradually lower maximal ETRs, steeper initial slopes of the P-I curves and, accordingly, lower light saturation points than upward-facing areas receiving higher light levels. Based on these trials, some evaluations are given as to the applicability of the Diving-PAM for photosynthetic measurements when monitoring similar corals.     

The mechanism of calcification and its relation to photosynthesis and respiration in the scleractinian coral <Emphasis Type="Italic"> Galaxea fascicularis</Emphasis>

The mechanism of calcification and its relation to photosynthesis and respiration were studied with Ca²⁺, pH and O₂ microsensors using the scleractinian coral Galaxea fascicularis. Gross photosynthesis (Pg), net photosynthesis (Pn) and dark respiration (DR) were measured on the surface of the coral. Light respiration (LR) was calculated from the difference between Pg and Pn. Pg was about seven times higher than Pn; thus, respiration consumes most of the O₂ produced by the algal symbiont's photosynthesis. The respiration rate in light was ca. 12 times higher than in the dark. The coupled Pg and LR caused an intense internal carbon and O₂ cycling. The resultant product of this cycle is metabolic energy (ATP). The measured ATP content was about 35% higher in light-incubated colonies than in dark-incubated ones. Direct measurements of Ca²⁺ and pH were made on the outer surface of the polyp, inside its coelenteron and under the calicoblastic layer. The effects on Ca²⁺ and pH dynamics of switching on and off the light were followed in these three compartments. Ca²⁺ concentrations decreased in light on the surface of the polyp and in the coelenteron. They increased when the light was switched off. The opposite effect was observed under the calicoblastic layer. In light, the level of Ca²⁺ was lower on the polyp surface than in the surrounding seawater, and even lower inside the coelenteron. The concentration of calcium under the calicoblastic layer was about 0.6 mM higher than in the surrounding seawater. Thus Ca²⁺ can diffuse from seawater to the coelenteron, but metabolic energy is needed for its transport across the calicoblastic layer to the skeleton. The pH under the calicoblastic layer was more alkaline compared with the polyp surface and inside the coelenteron. This rise in pH increased the supersaturation of aragonite from 3.2 in the dark to 25 in the light, and brought about more rapid precipitation of CaCO₃. When ruthenium red was added, Ca²⁺ and pH dynamics were inhibited under the calicoblastic layer. Ruthenium red is a specific inhibitor of Ca-ATPase. The results indicated that Ca-ATPase transports Ca²⁺ across the calicoblastic layer to the skeleton in exchange for H⁺. Addition of dichlorophenyldimethylurea completely inhibited photosynthesis. The calcium dynamics under the calicoblastic layer continued; however, the process was less regular. Initial rates were maintained. We conclude that light and not energy generation triggers calcium uptake; however, energy is also needed.