E任ect of Temperature on Photosynthesis and
Respiration in the Leaves of Di任erent Canopy
Layers within the Forest of Minamata Area
35
Tsukasa Kusumoto
Studies on dry matter production of the warm-temperate broad-leaved evergreen forest in
Minamata special-research-area of Japanese IBP-PT projects have been carried out since 1968.
●
In order to get some fundamental data concerning the forest productivity, the capacity of
●
photosynthesis and the activity of respiration of the leaves constituting the canopies on different
● ●
heights above the ground surface or at the different depths within the forest, and receiving the ● ●
in触ences from various experimental temperatures, were investigated throughout the years
from 1970 to 1973.
Hitherto, various issues occuring on a lot of species in a broad-leaved evergreen forest, i.e., ●
the seasonal activities of photosynthesis and respiration, the relationship between the capacities
●
of the photosynthesis and respiration and various temperatures, and the photosynthetic and
●
respiratory activities in the regions at di飴rent altitudes and latitudes, have been put under dis-
● ●cussion.6*11-13*17*19*22-33^7^9^5*5^^^^?^ On the other hand, many investigations of the effects
●
of temperature on the photosynthesis of coniferous and deciduous trees, at different seasons in various regions, have been carried out.5>8>9>19-21>27-38'40-54 63-68,7D in the reports, it was
gen-●
erally recognized that the three cardinal points on the photosynthesis*-temperature response
●
● ●
curve obtained in summer and in the regions of low altitude and low latitude showing macro-climatic conditions shi氏ed in the direction of higher temperature, and that the photosynthetic rates were ascertained to be larger than those in winter and in the regions of high altitude and
● high latitude.
As suggested by Larcher,32) concerning the leaf capacity within a forest canopy almost no
●
information has been supplied as yet. Therefore, the photosynthetic and respiratory capacities ●
of the leaves at different heights, growing under the diverse micro-climatic conditions within the
●
forest canopy, must be determined as exposed to the influences of various temperatures before the performance of the analysis of the process and structure of primary production in the forest.
36 Effect of Temperature on Photosynthesis
Materials and Methods
The structure and composition of broad-leaved evergreen forest in the JIBP-PT Minamata
special-research-area, and the phases of both the establishment for research and the regional●
environment, were reported by Hosokawa.10)
As the material plants were selected the dominant species in each stratum within the forest which were the same ones as adopted by Hozumi and Kirita.11} The dominant trees in upper layers were Castanopsis cuspidata and Quercus gilva, and they were of the height of 21m and about 50 years old. The dominant trees in lower layers of about 5m in height were Cleyera japonica, Euryajaponica and Machilusjaponica. In the used plants of the forest floor layer below lm in height, the low shrubs were ascertained to be Maesa japonica and Lasianthus japonicus,
and the herbs were Rubus Buergeri and Plagiogyria euphlebia (Pteridophyta).
The materials were collected in the sample plot of Quadrat 15 during the period from 1970 to 1971, and also in the place between P2 and P3 of permanent quadrat from 1972 to 1973. The leaves used for photosynthetic and respiratory measurements were chosen from twigs m various
●
heights, i.e., m C. cuspidata and Q. gilva, the heights above ground surface were 21, 19, 17, 15,
13, ll, 9, 5, 1 in meter(in Q. gilvaexcept llm in summer and 19m, 15m, 1lm, in winter). Counting
● from the youngest leaf of the first node of a twig apex, the detached mature leaves belonging to
from 2nd to 4th nodes were used for measurements. In Cl. japonica, E. japonica and M. japonica,
the heights of the trees out of which the material leaves were collected, were 5m and lm. In cases of Maesajaponica, L. japonicus, R. Buergeri and P. euphlebia composing the forest月oor layer,
●
the leaves of the plants from 0.2m to lm high were measured. The rates of gross photosynthesis and of dark respiration were determined with the infrared gas analyzers (Hitach Horiba Ltd. and Shimazu Ltd.) set in two cabins, which were constructed for the recording device and for sorting the collected materials and samples in the area. The measurements were carried out through all the seasons during the four from 1970 to 1973.
● ●
During the measurements the detached leaves in the assimilation chamber were illuminated
always at the saturated light intensity, above 40 Klux, with a National 500 W lamp. The light
intensity was measured with Toshiba Luxmeter No.5. The rate of air一月ow in the chamber was500 ml per minute, and on account of CO2 concentration of around 320 ppm in the atmosphere, the values obtained were corrected for 300 ppm CO2 concentration in calculation.
Tsukasa Kusumoto 37
which the chamber was placed was adjusted with a Taiyo Coolnit CL1 5 of therm-electric heating
and cooling apparatus. The leaf temperature was measured with a thermocouple attached to the reverse side of the leaf and the air temperature in the chamber was determined by a thermometer. The leaf area was measured with a planimeter and was indicated with the rate for one side of the
leaf.
Results
Gross photosynthesis
In the present paper, the values of gross photosynthesis of leaves at different heights were showed in the average for four years from 1970 to 1973. The gross photosynthesis-temperature response curves of leaves at different heights were illustrated in Figs. 1 and 2.
Among the curves obtained, those from the beginning of summer to late in autumn were ● ●
exhibited in a summer type, and after the first frost falling in the latter part of November, in a
● ● winter type. However, the different capacities were found in the leaves of various heights as the photosynthetic rates decreased with the lowering of the height of the leaf-attachment. The curves
●
in summer (summer type) and in winter (winter type) were the same as those in the results reported so far.22-26) The configurations of various curves shifting into a winter type from a summer type were observed in a certain period after the first frost falling, and so when the very cold seasons of January and February came the curves settled into the normal winter type. In the period in which turning of seasons occur, e.g., in case when the cold temperature was
●
changed into a warm one during the measurement, the curve of the early stage of the
measure-●
ment was similar to that of the con丘guration of cold condition, but they shi托ed to that of warm condition with the lapse of time. Therefore, the curves of irregular sigmoid shape or two low
●
peaks were observed in the above mentioned period.
The curves in summer. The curves in summer were determined during the period from the
●
latter part of May to the latter part of November. As shown in Fig. 1, the gross photosynthetic
curves of leaves at each height were entirely indicated by the optimum curves having a peak.●
Excepting the rate of21m which is smaller than that of 19m in C cuspidata and Q.gilva,the
maxi-mum rates of gross photosynthesis decreased in accordance with the lowering of the height of the●
leaves-attachment, as Hozumi et al. reported.1卜13) The leaves of the lower layer were observed to be the capacity of more shade leaf than those of the upper layer. The photosynthetic rate of the upper-most layer of crown with the height of 21m was smaller than that of the lower layer with the height of 19m and this fact corresponds with those ascertained by Hozumi and Kinta, and
38 Effect of Temperature on Photosynthesis
Pisek and Winkler.54) The curves of the forest floor plants were observed to be flat and low-hill-
●shaped as Koyama and Kawano18) appointed.
The ascertained optimum temperatures of the curves were illustrated in Fig 3. As shown in Figs. 1 and 3. in C. cuspidata the optimum temperatures of the upper-layer-leaves were 30--32-C and that of the leaves at the site of lm high was around 22-C, so that the optimum temperatures
of the loweトIayer-leaves seem to be apt to get lower. The trends of optimum temperatures in
Q. gilva were the same as those of C. cuspidata, and at the upper layers it was 28--30-C and at the
lowest height of lm it was about 27-C, which was higher than 23-C observed at the height of 5m.
In the trees of lower layer, such as Cl. japonica, E. japonica and M. japonica, the same pattern as
that of the upper layer was observed, too. However, the optimum temperature of M. japonica
was lower than those of two other species. The optimums of the forest floor plants such as Maesa
japonica, L. japonicus, R. Buergeri and P. euphlebia were around 24-C, which was lower than those
of the upper canopy layers. The turning-point of the optimum temperature between the upper and
●
the lower layers within the forest canopy was in the neighborhood of 13m in height. The similar trends may be recognized in aquatic plants.1"'4'i4-i-,56) From the above, it appears that the
●
special disposition, for example, a disposition that the optimum temperature of the upper layer is higher than that of the lower layer in the leaves within the forest canopy, is occasioned by the
●
effects of the micro-environmental conditions within the forest. The micro-climatic conditions within the forest of the area are reported in other papers.
The curves in winter. As shown in Fig. 2, in general the gross photosynthetic rates of winter, measured m the丘rst part of February, were smaller than those of summer, and the optimum and the maximum temperatures for photosynthesis shifted in the direction of low temperature as compared with those of summer. The results attained were the same as those of the author's reports22-26) for broad-leaved evergreen trees made so far. In the photosynthetic rates of leaves at different heights, the rate of 21m high in C. cuspidata was also smaller than those of 19m and 17m high as well as the curves in summer. However, in Q. gilva the rate of the one of 21m high was the maximum. From Fig. 2, the photosynthetic values in winter in C. cuspidata and Q. gilva were smaller than those m summer and those of the low-height-leaves in both species, decreased with the lowering of the height of leaf-attachment. The trends of photosynthetic values
●
in Cl. japonica, E.japonica and M. japonica were the same as those of the upper trees. Moreover,
the winter values in Maesa japonica, L. japonicus. R. Buergeri and P. euphlebia were smaller
than summer ones, respectively.Tsukasa Kusumoto 39
The optimum temperatures of leaves at different heights were arranged in Fig. 3. From such Fig. 3 as observed with a turning-point of around 13m in height, the optimum
●
temperatures of the upper layers were丘xed to be about 25-C> and those of the lower layers and the forest floor species were about 20-C, and found the tendency of dropping into lower
tern-●
perature in winter. But the dropping-range between the optimum temperatures of summer and
●
winter in Q. gilva was smaller than those in other species, and the optimum temperature of M.
japonica was the same both in summer and in winter and lower than those in other species, it being
● ●
20-C at 5m in height and about 17-C at lm in height, respectively. Summarizing the above results, in all the layers within the forest canopy the degrees of the dropping in winter-curves were small
● ●
and the decreasing values of photosynthesis became smaller with the lowering of the height of
● ● leaves. Also at each layer of all the species the optimums were of the same in winter.
The relation between the curves in summer and winter. On the three cardinal points of curves
Tablel Season Heights of leaf (m) Summer Speci es Rg I Rg T Rg Rg Castanopsis cuspidata Quercus gilva Cleyera japonica Eurya japonica ● Machilus japonica ● Maesa japonica Lasianthus japonicus Rubus Buergeri Plagiogyria euphlebia ;:53017.9-30. 827.516.5-29;.93 .230.;19.5-32 17-33 Season 8.3 32 21.5-35 7.1 28 17.5-31 8.1 30 19.5-31 6.4 29 17.5-31 Wi nter Castanopsis cuspidata Quercus gilva Cle vera japonica Eurya japonica ● Machilus japonica Maesa japonica● Lasian th us japonicus Rubus Buergeri Plagiogyria euphlebia 4.9 25 18.5-27 4.5 24 1 5-25 6.0 25 15-26.5 ,5:三芸.517-2 .515-2;4.42517.5-27
P The rates of gross photosynthesis at optimum temperatures (mg COzdm^hr"1). T The optimum temperatures ( C).
40 Effect of Temperature on Photosynthesis
of leaves at different heights in summer and winter, the minimum temperature both in sumriier and m winter was fixed to be around 5-C. Next, the optimum temperatures were different in summer and in winter, and in the upper layer it was about 30-C in summer and 250C in winter, and also in the forest floor plants it was 24-C in summer and about 20-C in winter. It was sup-posed that the ranges of falling into low temperature with the lowering of the leaf layer in the
●
optimum temperature of summer curves were larger than those of winter, as illustrated in Fig. 3 and Table 1. The maximum temperature through all the species was above 45-C in summer and above 35-C in winter. Likewise, the compensation-point of high temperature or the maximum temperature for net photosynthesis was observed to be about 40-C in summer and about 30 C in winter. Of the capacities of gross photosynthesis, excepting the height of 21m, the values of
winter in the upper layer were far smaller than those of summer and the decreasing degrees in ●
● ● winter became smaller with the lowering of the height of leaves. Therefore, concerning the
13 11 Rg Rg Rg Rg Rg 6.6 31 24-33.5 5.9 26 17.5-28 6.2 27.5 16-30 4.2 27 21-29 5.0 24 1 7-27 3.7 25.5 17-27 4.0 23 1 7-27 5.2 24 1 7-27 4.2 25.5 18-28 3.3 20 ll.5-22 2.4 22 16-25.5 3.2 27 21-30 3.4 23 1 7-26 3.8 22.5 17.5-28 2.8 17 10-19 3.0 24 1 7-27 2.7 25.5 15-26 4.1 25 16-26.5 2.7 21 1 5-24 4.2 25 1 7-26 3.4 25 1 8-27 3.8 22.5 15-25.5 ^o <n ^o i- h ● ● ● ● ● < N < N < r > m c M 20 1 4-23 20 1 6-24 20 1 4-24 20 14.5-26 20 1 5-21 1.9 20 17-23 2.3 21 1 6-24 3.3 20 1 6-23 3.1 20 15.5-26 1.8 17 14-22 1.8 20 14.5-22 2.0 20 1 5-22 2.7 20.5 14-25 2.4 19.5 1 1-22
Tsukasa Kusumoto 41 10 20 30 40 50 10 8 6 4 2 10 20 30 40 50 'wi^lざOォui 的isaq一uL等一｡qds等blOWH 8 6 4 ∧ 0 6 ▲1 L ^ J 0 8 6 4 2 0 10 20 30 40 50 _ : --:== 10 20 30 40 50 8 6 4 2 0 10 8 6 4 2 8 6 4 2 0 10 20 30 40 50 10 20 30 40 50 0 10 8 6 4 2 0 10 8 6 4 2 0 4 2 0 4 2 0 10 20 30 40 50 10 20 30 40 50
り均岳i ∵
･三二…二‥ =:=‥与‥ ≡= 10 20 30 40 50 Leaf temperature, CFig. 1. Temperature response curves of gross photosynthesis of leaves at different heights within the forest canopy in summer. The rates were measured on the detached mature leaves at the saturated light intensity and in the normal CO2-concentration of the atmosphere. 1 -4. Castanopsis cuspidata, 5-8. Quercus gilva, 9. Cleyera japonica, 10. Eurya japonica, 1 1. Machilus japonica, 12. Masea japonica (a), Lasianthus japonicus
Effect of Temperature on Photosynthesis ●● < o c * 10 20 30 40 ､50 j u / g i u p ＼ z q O S u i * s i S 9 i { ; u X s o ; o q d s s o j S j o 9 j B 出 8 6 4 2 n^K^ 10 20 30 40 50 10 8 6 4 2 0 10 8 6 4 2 0 10 20 30 40 50 5
■竺こ､
10 20 30 40 50 4 2 0 10 8 6 4 2 0 10 20 30 40 50 10 20 30 40 50 10 8 6 4 2 0 10 8 6 4 2 0j詰､
10 20 30 40 50 10 20 30 40 50 10 8 6 4 2 0 10 8 6 4 2 0 10 8 6 4 2 0 8 6 4 2 0 I 3s-10 20 30 40 50 13m :=^ 9m 10 20 30 40 50 9 5m lm 10 20 30 40 50 10 20 30 40 50 Leaf temperature, CFig. 2. Temperature response curves of gross photosynthesis of leaves at different heights within the forest canopy in winter. The rates were measured in the same conditions as in Fig. 1. 1 -4. Castanopsis cuspidata, 5- 7. Quercus gilva, 8. Cleyera japoncia, 9. Eurya japonica, 10. Machilus japonica, 1 1. Maesa japonica (a), La-sianthus japonicus(b), 1 2. Rubus Buergeri (a), Plagiogyria euphlebia(b).
Tsukasa Kusumoto 43
Winter
H牀 ･ T J Summer x O )0 x O O-C.cuspidata X -Q.gilva ACl.japonica o-'E.japonica ･x△oふ:::M.japonica Maesajaponica o L.japonicus ･R.Buergeri /?euphlebia･ 恥
■ × ○☆ ▲ 15 20 25 30 15 20 25 300ptimum temperature, ℃
35Fig. 3. Vertical changes in the optimum temperatures of the gross photosynthesis-temperature response curves in summer and in winter within the forest canopy.
44 Effect of Temperature on Photosynthesis
Rate of dark respiration, mgCCVdnr/hr.
●
○ ト■ N u Jゝ ul くつ トJ to c0 - もn OI ヽ1 くわ くつ ト to c0 - 01 0l ヽI ￠
欄!iliiiiヨi
5 10 15 20 25 30 35 40 45
Fig. 4. Temperature response curves of dark respiration of leaves at different height within the forest canopy in
●
summer. Also, those in winter were the same. 1 - 3. Castanopsis cuspidata, 4, 5. Quercus gilva, 6. Cleyera japonica (a, the curves of 5 m and 1 m indicate with one line), Machilusjaponica (5 m and 1 m), 7. Eurya japonica, 8. Maesajaponica(a)9 Lasianthusjaponicus(b), 9. Rubus Buergerifa), Plagiogyria euphlebia (b).
Tsukasa Kusumoto 45
di飴rence between the values in summer and winter it was large in the upper layers and small in
the lower layers, so that it seems that the productions of winter in the lower layers within the forest canopy are not decreased so much. The above results were summarized in Table 1.Dark respiration
The curves of the dark respiration of leaves at different heights were illustrated with the ●
average obtained during the period from 1970 to 1972 in Fig. 4. ●
The differences of the respiratory values in summer and winter were scarcely recognized, so ● ●
that only the curves of summer were shown. From the Fig. 4, the rates of the upper layer were ascertained to be larger than those of the lower layer at various temperatures and the rates de-creased with the lowering of the height of leaves and also each curve inde-creased exponentially,
●
when temperatures rose, through all the species.
Discussion
Hitherto, the investigations of the photosynthesis-temperature response curve have been carried out on many plants. It has been recognized obviously that of the many plants the
tempera-●
ture curves of evergreen plant exhibited two types; summer-type and winter-type, and that the photosynthetic capacities of summer were larger, and the cardinal points of summer were higher than those in winter.7>19>20>22>32>33>48) Thus, the experimental studies, in which a photosynthetic capacity which has been made approximately similar to that of summer is made to shi托into the winter capacity through the effect of cold temperature observable in natural conditions, were carried out with a lot of species of herbs, shrubs and trees under the arti丘cial conditions.58-60) The low temperature response-photosynthetic capacity of plants, exposed in a certain period under cold temperature condition, recovers that of warm temperature with the passing of time;
●
when treated under a warm temperature. The above photosynthetic activities have been re-cognized to be influenced by that of the term of "time factor".
●
During the experiment in the season in which the summer type is made to be changed into
●
the winter type after the frost falling, the capacity of early winter which was acclimated somewhat under natural cold conditions, was且uctuated by the length of the pre-acclimated time or the treated time of a moderate temperature, and the shi托ing curves of irregular sigmoid shape and
●
the curves showing two low peaks were observed. The above two curves seem to be particularly ●
46 Effect of Temperature on Photosynthesis
The photosynthesis-temperature response curves of the shaded leaves in the various canopy layers within the forest were determined in this paper, although the light intensity-response curves of the same forest canopy had been reported by Hozumi and Kirita. In the reports of Pisek et al.48>54) the optimums of the photosynthesis-temperature curves of both a sun-and a shade-leaf of coniferous trees, shifted in the direction of low temperature, when light intensities decreased, and the photosynthetic rate of shade leaf was always larger than a sun leaf under various light intensities owing to the rate calculated by the basis of leaf weight. The result
●
that the rate of21 m (semi-sun leaf) of top of the forest crown was a little less than that of 19 m (shade leaf) at a saturated light intensity, coincides with that ascertained by Pisek et al. But there is a difference in the fact that even under the saturated light intensity the optimum temperature into a low temperature with the lowering of the height of leaf and that the photosynthetic
●
rates of the lower layers are smaller than those of the upper layers both in the curve of summer
and in that of winter. Furthermore, the range of dropping into a 一ow temperature in summer-and ●
in winter-curves, were fixed to be about five degrees. Although the optimums of summer at each height触ctuated, it was le托the same in winter at each height through all the species. Therefore,
the falling degree of optimum temperature between the upper and the lower layers in summer seem to be larger than those in winter. Such phenomenon observable in the photosynthetic
capa-city of the leaves of di飴rent canopy layers indicates its adaptabilities to the micro-climatic
condition within the forest.
As for the dry matter productions, Pisek et al.51) a氏rmed that not the narrow apex of the curve but the range of temperature over 90% of the maximum net photosynthetic capcity was considered to be the temperature optimum for net photosynthetic yields. As shown in Table 1 the ranges of the present plants are fairly wide and around 10-C, from this standpoint it was assumed that when compared with the upper layer the range of curves of the forest floor plants was favor-able for the plant productions in order to be the very low and月at curves, as Koyama and Kawano
ascertained. Also, although the values of photosynthesis in winter were smaller than those of summer through all the layers, the decreasing rates were smaller in the lower layers and in the forest floor
●
plants than those in the upper layers. This capacity may be favorable for the plant production. As for the dark respiration, Larcher5>27>32>33) a氏rmed that the respiration of plants exposed
●
to the cold temperature indicated larger values after rewarming. In the Minamata area, on
ac-●
count of the absence of any excessive cold winter during the experimental years, and of the
con-●
ditions within the forest, the differences of the respiratory rates in summer and winter were left
Tsukasa Kusumoto 47
unobserved. However, if the respiratory activity for the cold temperature appears as
men-●
tioned above, the large rate of the one of21 m high at the top of the forest crown, exposed directly
to the open air condition of winter, may be noticeable in contrast with the small photosynthetic ●
rate. As a lot of reports suggested, the values of dark respiration of the leaves at different heights ●
within the forest canopy were decreased in accordance with the lowering of the height as well as in ●
case of the photosynthesis.
In a nutshell, the photosynthetic capacity and the respiratory activity of leaves at different ●
heights within the forest canopies are conducive to the dry matter production of the broad-●
leaved evergreen forest, respectively. It seems that the results are depending upon the effects of
the particular environmental conditions within the forest.
This study was carried out as a part of JIBP, Special Project Research "Studies on the
●
Dynamic Status of Biosphere", supported by the Grant in Aid of Scienti丘c Research of the
Ministry of Education.
Summary
In the study of the dry matter production of warm temperate broad-leaved evergreen forest in the JIBP-PT Minamata SpeciaトResearch-Area, the gross photosynthetic capacities and dark
respiratory activities of leaves, which constitute the canopy at di飴rent heights above ground
●surface within the forest, as influenced by various temperatures, were investigated and the values were indicated with the average during four years of the investigation.
●
The gross photosynthesis-temperature response curves exhibited the different types m summer and winter. The gross photosynthetic rates of leaves at different heights, excepting the small rate of leaves at the top of crown, decreased in accordance with the lowering of the height
●
of leaves. The rates of summer were larger than those of winter in all the heights and the decreas-ing degrees in summer at each height were larger than those in winter. In the
photosynthesis-●
temperature curves, the optimum and the maximum temperatures in summer were higher than in winter. The optimum temperatures in summer were about 30-C in the upper canopy layers and about 24-C in the lower layers, and so those in winter were 250C in the upper layers and about 20 C in the lower layers, respectively. The maximum temperatures in all the curves were above 45-C in summer and above 35-C in winter. The minimum temperature was the same both in summer and in winter and so 5-C. The optimum temperatures in summer and winter shifted in the
48 Effect of Temperature on Photosynthesis
direction of low temperature with the lowering of the canopy layers. The falling ranges of temper-●
ature were larger in the upper layers than those in the lower layers.
The dark respiratory activities were the same in summer and in winter on account of the ●
absence of any excessive cold winter during the investigation and the conditions within the forest ●
in the Minamata area. The dark respiratory rates of leaves at di飴rent heights decreased with the
●
● ●
lowering of the leaf layers in various temperatures and all of the respiration-temperature response
curves at different heights increase exponentially when temperatures rise.
It seems that the above results are depending upon the effects of particular environmental
conditions within the forest. A氏er all, the photosynthetic capacities and the respiratory activities ●
of leaves at different heights within the forest canopy are suitable for the dry matter production
oftheforest.
References
(*In Japanese with English summary. ** In Japanese)
1. Aruga, Y. (1965a) Ecological studies of photosynthesis and matter production of phytoplankton. I. Seasonal
changes in photosynthesis of natural phytoplankton. Bot. Mag. Tokyo 78, 280-288.
2. (1965b) Ibid. II. Photosynthesis of algae in relation to light intensity and temperature. Ibid. 78, 360
-365.
3. - (1966) Ibid. III. Relationship between chlorophyll amount in water and primary productivity. Ibid. 79, 20-27.
4. & Ichimura, S. (1968) Characteristics of photosynthesis of phytoplankton and primary production in the Kuroshio. Bull. Misaki Mar. Biol. Inst. Tokyo Univ. 12, 3-20.
5. Bauer, H., Huter, M. & Larcher, W. (1969) Der Einfluss und die Nachwirkung von Hitze-und Kaltestress auf den CO2-Gaswechsel von Tanne und Ahorn. Ber. deutch. bot. Ges. 82, 65-70.
6. , Larcher, W. & Walker, R. B. (1975) Influence of temperature stress on CO2-gas exhcange. In Photo-synthesis and Productivity in Different Environments, Intern. Biol. Progr. 3 (ed. J.P. Cooper), 557-586. Cam-bridge Univ. Press, CamCam-bridge.
7. Bliss, L.C. & Hadley, E.B. (1964) Photosynthesis and respiration of alpine lichens. Amer. J. Bot. 51, 870-847 8. Boysen Jensen, P. (1932) Die Stoffproduktion der Paanzen. Gustav Fischer, Jena, 108pp.
9. Furukawa, A. (1973) Carbon dioxide compensation points in popular plants. J. Jap. For Soc. 55, 95-99. 10. Hosokawa, T. (1969) An interim report of the ecosystem study of a warm-temperate evergreen borad-leaved
forest of the JIBP-PT Minamata Special Research Area in Japan. Malayan Forester 32, 405-41 3.
ll. Hozumi, K. & Kirita, H. (1970) Estimation of the rate of total photosynthesis in forest canopies. Bot. Mag. Tokyo 83, 144-151.
Tsukasa Kusumoto 49
IKJ
warm-temperate evergreen oak forest at Minamata (Japan). Photosynthetica 6, 1 58-1 68.
-, Nishioka, M., Nagano, M. & Kirita, H. (1973) Estimation of the total photosynthetic production by
forest canopies. Proc. East Asian Regional Seminar for the I.B.P. (ed. S. Mori & T. Kira), 45-56. Jap. Nat.
Com. I.B.P., Kyoto.
14. Ichimura, S. (1960) Photosynthesis pattern of natural phytoplankton relating to light intensity. Bot. Mag.
Tokyo 73, 458⊥467.
1 5. (1961) On the spatial difference of the primary production in the lake and its relation to environmental
factors. Ibid. 74, 6-13.
16. Ikusima, I. (1965) Ecological studies on the productivity of aquatic plant communities. I. Measurement of
photosynthetic activity. Ibid. 78, 202-21 1.
17. Kan, M., Saito,H. & Shidei, T.(1965)Studies of the productivity of evergreen broad-levaed forest. Bull. Kyoto Univ. For. 37, 55-75*.
18. Koyama, H. & Kawano, S. (1973) Biosystematic studies of Mainanthemum (Liliaceae-Polygonatae). VII.1
Photosynthetic behaviour of M. dilataum under changing temperate woodland enivornments and its biologi-cal implications. Bot. Mag. Tokyo 86, 89-101.
19. Kramer, P.J. & Kozlowski, T.T. (1960) Physiology of trees. McGraw-Hill, New York, Tronto & London.
642pp.
20. Kuroiwa, S. (1960) Ecological and physiological studies on the vegetation of Mt. Shimagare. IV. Some physiological functions concerning matter production in young Abies trees. Bot. Mag. Tokyo 73, 133-141.
●
21. & Monsi, M. (1956) On the relation of light and temperature to the structure of plant community. J. Agr. Meteorol. 12, 41｣¥7 *
22. Kusumoto, T. (1957a) Physiological and ecological studies on the plant production in plant communities.
23.
24.
25.
26.
3. Ecological consideration of the temperature-photosynthesis curves of evergreen broad-leaved trees. Jap.
J. Ecol. 7, 126-130*.
(1957b) Ibid. 4 Ecological studies on the apparent photosynthesis curves of evergreen broad-leaved
trees. Bot. Mag. Tokyo 70, 299-304.
(1957c) Ibid. 5 On the photosynthesis of broad-leaved evergreen trees in Amami Oshima Island.
Bull. Fac. Educ. Kagoshima Univ. 9, 21-25*.
(1963) Ibid. 8. The activities of photosynthesis and respiration in the broad-leaved evergreen trees
growing in the littoral region or in the highland in Kagoshima Prefecture. Ibid. 15, 64-73.*
(1961) An ecological analysis of the distribution of broad-leaved evergreen trees, based on the dry
matter production. Jap. J. Bot. 17, 307-331.
27. Larcher, W. (1961a) Jahresgang des Assimilations-und Respirationsvermogens von Olea europaea L. ssp.
28.
sativa Ho ff. et Link., Quercus ilex L. und Quercus pubescens Willd aus dem nordlichen Gardaseegebiet.
Planta 56, 575-606.
som-50 Effect of Temperature on Photosynthesis 29. 30. 31. 32. 33. mergriinen i t. Ibid. 56, 607-617.
(1963a) Die Eignung abgeschnittener Zweige und Blatter zur Bestimmung des Assimilationsver-mogens. Ibid. 60, 1-18.
蝣(1 963b) Die Leistungsfahigkeit der CO2-Assimilation hoerer Pflanzen unter
Laboratoriumsbedingun-gen und am natむIichen Standort. Mitt. Flor.-soz. Arbeitsgem. Arge. 10, 20-33.
(1969a) Die Bedeutung des Faktors "Zeit" fiir die photosynthetische Stoffproduktion. Ber. deut. bot. Ges. 82, 7ト80.
- (1969b) The effect of environmental and physiological variables on the carbon dioxide gas exchange of trees. Photosynthetica 3, 1 67-198.
- (1975) Physiological Plant Ecology. Spring Verlag, Berlin, Heidelberg & New York. 252pp.
34. Ldhr, E. (1969) Respirationsintensitat in Stammen, Zweigen und Blattern von Laubbaumen in torpischen
Regenwald und in temperieten Waldern. Physiol. Plant. 23, 86-93.
35. Lundegardh, H. (1957) Klima und Boden in ihrer Wirkung auf das Pflanzenleben. 5 Au札Gustav Fischer,
Jena. 598pp.
36. Maruyama. K. (1971) Effect of altitude on dry matter production of primeval Japanese beech forest com-mumties in Naeba Mountains. Mem. Fac. Agr. Niigata Univ. 9, 87-171.
37. & Teshima, T. (1972) Ecological studies on natural beech forest. 29. Comparative studies on the rates
38.
of photosynthesis among the di飴rent Japanese broad-leaved tree species (I). Niigata Norin Kenkyu 24,
31｣12*
- & Yamada, M. (1968) Ibid. 16. Seasonal course on apparent photosynthesis and respiration rate in
detached leaves of Japanese beech at different localities. Bull. Niigata Univ. For. 3, 17-26.*
39. Miyata, I. (1970) Analysis of the life table of leaves of Castanopsis cuspidata. I. Seasonal changes of the rate
of photosynthesis in single leaves. Studies on the biological production of warm-temperate evergreen
broad-●
leaf forests, Progress report for 1969 (ed. T.Hosokawa), 3ト34.*
40, Mooney, H.A. & West, M. (1964) Photosynthetic acclimation of plants of diverse origin. Amer. J. Bot. 51,
825-827.
41. - & Harrison, A.T. (1970) The influence of conditioning temperature on subsequent temperature-related photosynthetic capacity in higher plants. In Prediction and Measurement of Photosynthetic Produc-tivity, Proc. IBP/PP Tech. Meet., Trebon, 1969, 41 1-417. Centre for Agricultural Publishing and Documen-tation, Wageningen.
●
42. Murata, Y., Iyama, J. & Homma, T. (1965) Studies on the photosynthesis of forage crops. 5. Influence of
air-temperature upon the photosynthesis and respiration of alfalfa and several southern type forage crops. Proc. Crop. Sci. Soc. Jap. 34, 154-158.
43. Negisi, K. (1966) Photosynthesis, respiration and growth in 1-year-old seedlings of Pinus densiflora,
Crypto-44.
meriajaponica and Chamaecyparis obtusa. Bull. Tokyo Univ. For. 62, 1-1 15.
Tsukasa Kusumoto 51
densiflora Sieb. et Zucc.), Sugi (Cryptomeria japonica D. Don.) and Hinoki (Chamaecyparis obtusa Sieb. et Zucc), J. Jap, For. Soc. 43, 336-343.
45. Nomoto, N. (1964) Primary productivity of beech forest in Japan. Jap. J. Bot. 18, 385-421.
46･ -, Kasanaga, H. & Monsi, M. (1959) Dry matter production by Chamaecyparis pisifera in winter. Bot. Mag. Tokyo 72, 450-455.
47. Pisek, A. (1960a) Paanzen der Arktis und des Hochgebirges. Handb. d. Pflanzenphysiol. (ed. W.Ruhland)
50. 51. 52. 53. 54. V/2, 376-414.
(1 960b) Immergr屯ne P且anzen (Einschliessliche Coniferen). Ibid. V/2, 41 5-459.
- (1963) Zur Kenntnis der Temperaturabhangigkeit der Netto-Assimilation von Samenpflanzen. Mitt. Flor.-soz. Arbeitsgem. Arge. 10, 34-41.
- & Kennitzer, R. (1968) Der EinRuss von Frost auf die Photosynthese der Weisstanne. Flora B 157,
314-326.
-, Larcher, W., Moser & Pack, I. (19691 Kardinale Temperaturbereiche der Photosynthese und Grenz-temperaturen des Lebens der Blatter verschiedener Spermatophyten. III. Temperaturabhangingkeit und
●
optimaler Temperaturbereich der Netto-Photosynthese. Flora B 1 58, 608-630.
-, Pack, I. & Unterholzer, R. (19681 Ibid. II. Temperatur maximum der Nettophotosynthese und Hitzresistanz der Blatter. Ibid. B 158, 1 10-128.
& Unterholzer, R. (1967) Ibid. I. Temperaturminimum der Netto-Assimilation, Gefrier-und Frostschadensbereiche der Blatter. Ibid. B 1 57, 239-264.
& Winkler, E. (1959) Licht-und Temperaturabhangigkeit der CO2-Assimilation von Fichte (Picea excelsa Link.), Zirbe (Pinus cembra L.) und Sonnenblume (Helianthus annuus L.). Planta 53, 532-550.
55. Saeki, T. & Nomoto, N. (1958) On the seasonal change of photosynthetic activity of some deciduous and evergreen broad-leaf trees. Bot. Mag. Tokyo 71, 235-241.
56. Saitoh, M., Narita, K. & Isikawa, S. (1970) Photosynthetic nature of some aquatic plants in relation to temperature. Ibid. 83, 10-12.
57. Sanai, H., Nakayama, A., Kano, T. & Sakai, S. (1967) Studies on the changes of internal constituents with the growth in young tea plant. Bull. Tea Res. St., Ministry ofAgr. & For. 4, 1-33.*
58. Sawada, S. (1970) An ecophysiological analysis of the difference between the growth rates of young wheat seedlings grown in various seasons. J. Fac. Sci. Tokyo Univ. Ser. Ill, 10, 233-263.
59. - (1973) Net assimilation rates of wheat plants grown in two fields with different temperature condi-tions during winter. Jap. J. Ecol. 23, 234-250.
60. & Miyachi, S. (1974) Effects of growth temperature on photosynthetic carbon metabolism in green plants. I. Photosynthetic activities of various plants acclimatized to varied temperature. Plant & Cell Physiol.
15,11ト120.
61. Strain, B. R. & Chase, V. C. (1966) Effect of past and prevailing temperatures on the carbon dioxide exchange
52
Effect of Temperature on Photosynthesis
62. Tadaki, Y. (1968) Studies on the production structure of forest (14). The third report on the primary produc-tion of a young stand of Castanopsis cuspidata. J. Jap. For. Soc. 50, 60-65.
63. Tranquillini, W. (1959a) Die Stoffproduktion der Zirbe (Pinus cembra L.) an der Waldgrenze wahrend eines Jahres. I. Standortklima und CO2-Assimilation. Planta 54, 107-129.
64. - (1959b) Ibid. II. Zuwachs und CO2-Bilanz. Ibid. 54, 130-151.
65. & Schutz, W. (19701日ber die Rindenatmung einiger Baume and der Waldgrenze. Centralbl. f.
gesamte Forstwesen 87, 42-60.
66. Ungerson, J. & Scherdin, G. (1965) Untersuchungen屯ber die photosynthese und Atmung unter natiirlichen
Bedingungen wahrend des Winterhalbjahres bei Pinus silvestris, Picea excelsa und Juniperus dommunis. Planta 67, 136-167.
67. - & (1968) Jahresgang von Photosynthese und Atmung unter natiirlichen Bedingungen bei Pinus silvestris and ihrer Nordgrenze in der Subarktis. Flora B 157, 391-434.
68. Winkler, E. (1961) Assimilationsvermogen, Atmung und Ertrage der Kartoffelsorten Oberarnbacher Friihe, Planet, Lori und Agnes im Tal (610m) und an der Waldgrenze bei Innsbruck und Vent (1880m bzw. 2014m). Ibid. 151, 621-662.
69. Yoda, K. (1967) Comparative ecological studies on three main types of forest vegetation in Thailand. III. community respiration. Nature & Life in SE Asia 5, 83-148.
70. Yoda, K., Shinozaki, K., Ogawa, H., Hozumi, K. & Kira, T. (1965) Estimation of the total amount ofres-piration in woody organs of trees and forest communities. J. Biol. Osaka City Univ. 16, 15-26.
71. Zelawski, W. & Kucharska, K. (1967) Winter depression of photosynthetic activity in seedlings of scots pine (Pinus silvestris L.). Photosynthetica 1, 207-213.
