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CHILLING SENSITIVE
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FIG. 1. Arrhenius plots of succinate oxidation by plant mitochondria. Each plot showing state 3 (0) and state 4 (0) respiration represents
data from three or more mitochondrial preparations. The log values were adjusted by a factor to a common value at 25 C in order to compensate
for differences in the rates (nanomoles of 02 per min per mg of protein) between the different preparations.
Downloaded from www.plantphysiol.org on October 27, 2014 - Published by www.plant.org
Copyright © 1970 American Society of Plant Biologists. All rights reserved.
388 LYONS AND RAISON Plant Physiol. Vol. 45, 1970
(22) for mitochondria from pears, the RC ratios shown in Table is not affected directly by low temperatures although the rate of
I fell within the range usually observed for succinate oxidation phosphorylation would be reduced concomitant with reduction
by plant mitochondria (1, 3, 5, 23, 26, 27, 30) and by rat liver in oxidative activity.
mitochondria (6). Similarly, the ADP:O ratios, although ex- The relationship between respiratory activity and chilling
hibiting some variability, show no consistent pattern which can injury has been investigated as a postharvest problem with
be related to temperature effects and approach the theoretical intact tissues (cf References 2, 9, 19, 28), tissue slices (10, 17,
value of 2 for succinate oxidation. Discontinuities in the Ar- 24), excised roots (29), and mitochondria (12, 16). In relating
rhenius plot for succinate oxidation and for the ADP-ATP and these studies on respiratory behavior and chilling injury, it is
Pi-ATP exchange reactions catalyzed by rat liver mitochondria essential to distinguish between the primary response to chilling
have been reported (7). Each of the reactions measured show a temperatures, and subsequent responses associated with altered
change in the temperature coefficient at approximately 17 C with metabolism. For example, if the data of Eaks and Morris (2)
a 2-fold increase in the Qlo value below the critical temperature. for chilling sensitive cucumber fruit are presented as an Ar-
We have measured the rate of succinate oxidation by rat liver rhenius plot of the respiration rates at 24 hr and 5 and 10 days
mitochondria and found a discontinuity in the Arrhenius plot (Fig. 2) after being placed at the test temperatures, the differ-
between 23 and 25 C for both state 3 and state 4 respiration. ences between the initial effect of temperature on respiration and
those changes in respiration which occur after injury are readily
DISCUSSION apparent. At 24 hr (the earliest measurement reported), the
The results clearly demonstrate that the respiration of mito- respiration rates diminish with a Qlo value of 1.7 from 30 to
13 C. A change occurs between 10 and 13 C. Below 10 C there
chondria from chilling sensitive plant tissue, in contrast to the is an increase in the rate of reduction of respiration as the tem-
mitochondria from resistant tissue, is significantly reduced at perature is decreased further. The Qlo value rose to 2.9. Eaks
temperatures below 10 C, indicating that the immediate response and Morris (2) reported visible injury between 24 hr and 5 days
of sensitive tissue to chilling is a depression of respiratory activ- in the fruit held at 0 and 5 C, and this injury was reflected by
ity. This depression in activity observed in each of the sensitive the increased respiration rates at these temperatures. By 10
tissues below a critical temperature is consistent with the hy- days in storage, injury and increased respiration had occurred at
pothesis that reduced temperatures affect some physical property all temperatures at or below 10 C, causing the slope of the Ar-
of the mitochondrial membranes, e.g., membrane lipids (13, 14, rhenius plot in this temperature region to be elevated to the
21). At the same time, the data show that phosphorylative effi- slope exhibited at the higher temperatures (Fig. 2). An Arrhenius
ciency of mitochondria from sensitive as well as resistant tissues plot of the data for intact potato tubers (19) shows a linear slope
Table I. Respiratory Conitrol antd Oxidative Phosphorylation over the entire temperature range from 25 to 0.5 C (Fig. 2).
for Succinate Oxidationl by Plaint Mitocholzdria While the rate of respiration gradually decreases with time in
Data for respiratory control (RC) ratios and phosphorylation
storage (as does the respiration of cucumbers at nonchilling
(ADP:O ratios) are averages + the standard error of two to five
temperatures, Fig. 2), there is no deviation in the linearity of the
cycles from three to five experiments for each of the tissues shown slope. With the use of respiration data obtained from intact
in Figure 1. Procedures are described in "Materials and Methods."
tissues as well as the mitochondrial data (Fig. 1), it is evident
that a very early response to temperatures below the critical
Respiratory Control ADP:O point for injury is a depression in respiratory activity. This
Temp depression in respiration is not observed with mitochondria
Resistant Sensitive Resistant Sensitive (Fig. 1) or plant tissues (Fig. 2) resistant to chilling injury.
c
Impairment of mitochondrial phosphorylation is not an
immediate result of chilling (Table I), but probably follows after
0-4.9 3.4 I 1.1 2.4 + 0.3 1.35 + 0.12 1.44 i 0.10 the tissues have been injured for some time period. Schichi and
5-9.9 2.8 0.3 2.2 ± 0.3 1.40 ± 0.07 1.67 + 0.15
10-25 2.5 i 0.3 2.3 ±4- 0.2 1.76 + 0.13 1.47 + 0.18 Uritani (24) demonstrated that sweet potato disks did not in-
crease respiration in response to dinitrophenol after the roots
POTATO TUBERS
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33 4 3s 3' 35 36
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FIG. 2. Arrhenius plots of respiration data for intact cucumber fruit and potato tubers. Data for cucumber fruit calculated from Eaks and Morris
(2) and for potato tubers from Platenius (18). The different slopes represent different time periods (as indicated) after tissues were placed in the
temperature treatments.