Influence of depth and transparency

Once established, aquatic macrophytes have a positive effect on the transparency of water through several buffer mechanisms (Stephen et al. 1998). Furthermore, the presence of charophytes has been associated with the maintenance of clear water, and changes from a state of clear to turbid water have been associated with the eutrophication of the environment (e. g. Blindow et al. 1993, Kufel & Kufel 1997).

Steinman et al. (1997) studied the influence of water depth and transparency on the charophyte biomass distribution in the southern end of the subtropical Lake Okeechobee, U. S.A. Their first survey (August 1994) was conducted on 47 stations within the 3-Pole Bay. Subsequent surveys (November 1994-December 1996) were conducted on a monthly or bimonthly basis on 7 stations. According to the authors, the distribution and abundance of Chara population in the lake showed a marked seasonal phenology, although there were notable differences in biomass among the years and stations. Chara plants were observed only in August, September and October, and in 1996 also in November. Also, biomass never exceeded 20 g AFDM m-2 and declined significantly from 1994 to 1996. The charophyte biomass was inversely related to the water depth and positively related to the Secchi disc depth, suggesting that irradiance strongly influenced the charophyte distribution in the lake, a hypothesis that was confirmed by data they collected from photosynthetic measurements and phytosynthesis-irradiance curves (Steinman et al. 1997).

The role of charophytes in increasing the water transparency was also studied by N5ges et al. (2003). Under the frame of the EC project ECOFRAME, last authors worked out the water quality criteria for two shallow lakes of the Vooremaa landscape protection area, Central Estonia. Lake Prossa is a macrophyte-dominated system with an area of 33 ha and a mean depth of 2.2 m. Most of its bottom is covered by a thick mat of charophytes all year round. Lake Kaiavere is located 10 km far from Lake Prossa, is much larger (250 ha, mean depth 2.8 m) and is phytoplankton-dominated. Nevertheless, the nutrient dynamics was very similar in the two lakes (Figure 4). The first vernal phytoplankton peak was expressed in reduced Secchi depth in both lakes. After that peak, the water became clear in Lake Prossa, but remained turbid in Lake Kaiavere. Towards fall, the individual mean weight of zooplankton decreased in Lake Prossa, the Chara-lake, but remained smaller than in the plankton — dominated one (Lake Kaiavere) (Figure 5). Therefore, zooplankton grazing would initiate the clear-water phase in the Chara-lake, but other factors were needed for its maintenance. Another factor that showed a clear difference between the two lakes was the carbonate alkalinity that was rather stable or even increased during the spring in the phytoplankton — dominated lake, while it decreased by nearly 50% between April and July in the Chara-lake. The reduced sediment resuspension and the possible allellopathic influence of charophytes on phytoplankton remain the main explanations for the maintenance of the extensive clear­water period in the Chara-lake.

Blindow & Schutte (2007) worked with material from fresh and brackish water in Sweden and found out that both turbidity and salinity acted as stress factors on Chara aspera. According to the last authors, in clearwater lakes the species can occur in high densities and reach deep water, where the ability to hibernate as a green plant together with shoot elongation may further extend the lower depth limit. In turbid lakes, the plants can still form dense mats, but are restricted to shallow water due to the poor light availability, although shoot elongation may allow a certain extension of the depth range (Figure 6).

Figure 4. Seasonal dynamics of some chemical and biological features in lakes Prossa (Chara-lake) and Kaiavere (plankton-lake). A — total nitrogen, B — total phosphorus, C — dissolved organic nitrogen, D — soluble reactive phosphorus, E — zooplankton/phytoplankton biomass ratio, G — zooplankton mean individual weight, H — hydrocarbonate concentration, I — pH, and J — calcite saturation level (Noges et al. 2003).

Figure 5. Left: total length of Chara aspera (mean values + S. E., n = 5), determined at all sites, depth ranges and sampling occasions. Above freshwater sites, bellow brackish water sites. K — Lake Krankesjon, B — Lake Borringesjon, S — Lake Storacksen, E — Edenryd bay, H — Hollviken bay, R — Redensee bay. I and II — first and second sampling occasions, respectively. Right: fresh weight:total length ratio of Chara aspera determined at all sites, depth ranges and sampling occasions (Blindow & Schutte 2007).

A field study conducted from July 2003 to May 2005 in the Myall Lake, a brackish shallow lake in New South Wales, Australia, revealed that Chara fibrosa var. fibrosa and Nitella hyalina occurred in areas of the entire lake that were deeper than 50 cm. Also, more fresh shoots were obtained during the winter (water temperature 13-16°C), thus suggesting that winter may be their preferred growing season. Their biomass varied from 0 to 321 g DW m-2, their maximum biomass being displayed between 1 and 2.5 m depth (Asaeda et al. 2007). These authors also observed that charophyte’s shoots were longer in deeper waters, varying from c. 30 cm at 1 m depth to 60-90 cm between 2 and 4 m depth. Plants growing in shallow depths had shorter internodes implying a shorter life cycle of shoots. Also, nodal spacing was relatively regular in contrast to its deeper water counterparts although spacing tended to increase at locations farther from the apex (Figure 7). Finally, numbers of oospore and antheridia were higher in shallower water reaching their maximum at around 80 cm.

Chambers & Kalff (1985) used original data from eight lakes in southern Quebec, Canada and literature data from other lakes throughout the World to predict the maximum depth of charophytes colonization and the irradiance over the growing season at the maximum depth of colonization, concluding that the depth distribution of the aquatic macrophyte communities is quantitatively related to Secchi depth. According to regression models proposed in Chambers & Kalff (1985), natural distribution of aquatic macrophytes is restricted to depths of less than 12 m, whereas charophytes can colonize to great depths and up to a predicted 42 m in the very clearest lakes (Secchi depth 28 m).

Figure 6. Seasonal variation of (a and b) Chara fibrosa and (c and d) Nitella hyalina biomass at location deeper than and shallower than 1 m. "X" markers denote individual measurements, the thick solid line represents monthly means, and short flat bars indicate standard deviations (mean + 1 S. D.) (Asaeda et al. 2007).