Growth responses of native mosses at low temperature
At the low temperatures of 15/5°C or 5°C, all mosses showed a significant decrease in block area regardless of temperature, which dropped down to 50–70% of the initial area at first one month (
Fig. 2). After that, there was not much difference in moss block area for further one month. This suggests that when the minimum temperature drop to 5°C, mosses are stunted in growth or discolored, showing a rapid decrease of area, but after that they showed almost no growth. Mosses respond sensitively to environmental changes, and thus their growth is easily discontinued, and they enter dormancy if the weather becomes too dry or cold. Stunted growth of mosses at low temperature varied among species.
Etodon luridus and
Hypnum erectiusculum showed the least area decrease (
Fig. 2B, 2I), whereas
Marchantia polymorpha, Bachythecium plumosum, and
Hypnum plumaeforme showed the most area decrease (
Fig. 2C, 2F, 2H). The other five types were similar.
After low temperature treatment, the plant growth temperature was set to 20°C and the recovery degree of mosses was examined for 6 weeks. It was found that moss block area rather decreased in the first 2 weeks, showing approximately 20% decrease than the low temperature state (
Fig. 2). After that, moss block area gradually increased over time. Moss block area rather decreased in the first 2 weeks even when growth temperature increased to 20°C, which is the optimum growth temperature of moss. It seems likely that any symptom of low temperature injury was not shown, because most mosses easily stop growth or begin dormancy at 5°C, but the symptoms of injury appeared on the surface after metabolic activities started at optimum growth temperature. The recovery of mosses at optimum growth temperature varied among moss species.
Etodon luridus,
Myuroclada maximowiczii, and
Hypnum erectiusculum showed the most recovery in moss block area compared to other mosses (
Fig. 2B, 2E, 2I). They showed 70–80% recovery of the initial area at 6 weeks at room temperature. The mosses having the biggest injury at the low temperature and the least recovery at room temperature were
Atrichum rundulatum,
Marchantia polymorpha,
Thuidium cymbifolium, and
Hypnum plumaeforme, which were merely 50% of the initial area in 6
th week at 20°C (
Fig. 2A, 2C, 2D, 2H).
Changes in moss color after low temperature and at room temperature
When mosses are used in indoor or outdoor landscaping, they look fresh in green and thus green mosses have high utilization value. However, mosses easily enter dormancy when the temperature drops in winter and the color turns brown, thus likely to lose ornamental value. Therefore, the ability of moss quickly to recover at room temperature is very important characteristics.
Table 1 and
Fig. 3 show the changes in moss color after low temperature treatment and at optimum growth temperature. When mosses are treated at low temperatures of 15/5°C (16/8h, day/night) or 5°C (24h), red, green and blue values all decreased greatly. All color values that decreased at 4 weeks of low temperature treatment regardless of the temperature remained almost the same by 8 weeks as well. After the temperature was transferred to optimum temperature later, the recovery of green varied among moss species. Red and blue did not show any correlation with moss block area or growth state, and only green showed a correlation with moss block area or growth state, which is why we will explain changes in green values here. In fact, it is more important to maintain greenness in reality as well.
At low temperature of 15/5°C, Etodon luridus showed 87% of initial value in green color, and 90% at room temperature. Hypnum plumaeforme also had 85% of the initial greenness at low temperature, and increased to 98% at room temperature, showing that it is well maintaining greenness despite the reduced area by low temperature and revealing more greenness at room temperature. Bachythecium plumosum decreased in greenness to 70% of initial value at low temperature, but recovered up to 90% at room temperature. Therefore, these three mosses are strong against low temperature and quickly recover at room temperature after low temperature treatment.
On the other hand, the species that showed the most remarkable decrease in greenness at low temperature were Atrichum undulatum and Marchantia polymorpha, which had their greenness drop to below 50% of the initial greenness. This suggests that even when the temperature setting was changed to room temperature, the greenness was not recovered and thus completely withered, or showed barely any recovery at all. Thuidium cymbifolium damaged by low temperature, failed to recover even at room temperature, and the greenness gradually decreased from 60% to 40%. The other three species such as Myuroclada maximowiczii, Plagiomnium cuspidatum, and Hypnum erectiusculum showed a moderate decrease in greenness at low temperature and moderate recovery of greenness at room temperature.
Treatment at 5°C (24 h) also showed similar results with 15/5°C temperature treatment. Etodon luridus and Hypnum plumaeforme showed the least decrease in greenness at low temperature, and showed high greenness at room temperature. Only Bachythecium plumosum showed little change in greenness at 15/5°C but showed decrease of up to 60% of the initial value, while also rarely showing any recovery. On the other hand, Atrichum undulatum, Marchantia polymorpha, and Plagiomnium cuspidatum showed decrease in greenness by 50% of the initial value, and there was barely any change in greenness even at room temperature. Thuidium cymbifolium, like at 15/5°C, showed continuous decrease in greenness even at room temperature.
This tendency was found more precisely in actual photographs (
Fig. 3).
Etodon luridus maintained greenness without any injury at the two low temperatures and retained vivid green at room temperature, showing greatest tolerance to cold.
Bachythecium plumosum did not show severe injury at low temperature of 15/5°C, and also seemed healthy at room temperature.
Hypnum plumaeforme showed decrease in greenness due to low temperature, but the green was vivid on living mosses, and showed more vivid greenness due to room temperature.
In summary of the moss block area, greenness and actual images,
Etodon luridus that showed the least change in moss block area and also in green value (
Table 1,
Fig 3).
Bachythecium plumosum showed a great decrease in moss block area at 15/5°Ct, but maintained the greenness without much decrease due to low temperature. These results indicated that these 2 species are relatively strong against low temperature.
Hypnum plumaeforme showed a great decrease in moss area due to low temperature, but living mosses did not turn brown but showed vivid green, thereby maintaining ornamental value.
Hypnum erectiusculum shown a little decrease in moss block area due to low temperature as well as high resilience, showed a significant decrease in greenness, indicating that it is not strong against low temperature.
On the other hand, Atrichum undulatum, Marchantia polymorpha, and Thuidium cymbifolium which showed a great decrease in moss area at low temperature and did not show a great increase in moss area at recovery temperature, had significantly lower greenness than other species, thereby proving to be sensitive to low temperature. The other species such as Myuroclada maximowiczii, Plagiomnium cuspidatum and Hypnum erectiusculum showed moderate responses low temperature.
Furness and Grime (1982) also examined growth responses to temperature by cultivating 40 species of bryophytes in various temperature ranges, and found out that most mosses showed the best growth at 15–25°C, but only
Dicranella palustris and
Racomitrium lanuginosum grew well at the low temperature of 12–13°C. More than half of the species showed a decrease in growth rates to below 50% at 5°C, indicating that responses to low temperature varied among species.
Atrichum undulatum has leaves that curl up when dry and uncurled when there is water, thereby having a great ornamental value (
Choi, 1980), but it is sensitive to low temperature.
Myuroclada maximowiczii (G.G. Borshch.) Steere & W.B. Schofield has round leaves and smooth stems and branches, thus suitable for indoor landscaping, but they stop growing and turn brown when the temperature drops to below 5°C in winter. Indoor or outdoor landscaping of building using mosses can reduce heating and cooling costs and alleviate the phenomenon of urban heat island (
Lee et al., 2005), and thus use of mosses is expected to increase for roof or wall planting. However, when using mosses, it is important to choose the ones with high resistance to low temperature as well as resilience.