Lc. Van Ginkel et al., A CO2-flux mechanism operating via pH-polarity in Hydrilla verticillata leaves with C-3 and C-4 photosynthesis, PHOTOSYN R, 68(1), 2001, pp. 81-88
The aquatic angiosperm Hydrilla verticillata lacks Kranz anatomy, but has a
n inducible, C-4-based, CO2 concentrating mechanism (CCM) that concentrates
CO2 in the chloroplasts. Both C-3 and C-4 Hydrilla leaves showed light-dep
endent pH polarity that was suppressed by high dissolved inorganic carbon (
DIC). At low DIC (0.25 mol m(-)3), pH values in the unstirred water layer o
n the abaxial and adaxial sides of the leaf were 4.2 and10.3, respectively.
Abaxial apoplastic acidification served as a CO2 flux mechanism (CFM), mak
ing HCO3- available for photosynthesis by conversion to CO2. DIC at 10 mol
m(-3) completely suppressed acidification and alkalization. The data, along
with previous results, indicated that inhibition was specific to DIC, and
not a buffer effect. Acidification and alkalization did not necessarily sho
w 1:1 stoichiometry; their kinetics for the apolar induction phase differed
, and alkalization was less inhibited by 2.5 mol m(-3) DIC. At low irradian
ce (50 mu mol photons m(-2) s(-1)), where CCM activity in C-4 leaves is min
imized, both leaf types had similar DIC inhibition of pH polarity. However,
as irradiance increased, DIC inhibition of C-3 leaves decreased. In C-4 le
aves the CFM and CCM seemed to compete for photosynthetic ATP and/or reduci
ng power. The CFM may require less, as at low irradiance it still operated
maximally, if [DIC] was low. Iodoacetamide (IA), which inhibits CO2 fixatio
n in Hydrilla, also suppressed acidification and alkalization, especially i
n C-4 leaves. IA does not inhibit the C-4 CCM, which suggests that the CFM
and CCM can operate independently. It has been hypothesized that irradiance
and DIC regulate pH polarity by altering the chloroplastic [DIC], which ef
fects the chloroplast redox state and subsequently redox regulation of a pl
asma-membrane H+-ATPase. The results lend partial support to a down-regulat
ory role for high chloroplastic [DIC], but do not exclude other sites of DI
C action. IA inhibition of pH polarity seems inconsistent with the chloropl
ast NADPH/NADP(+) ratio being the redox transducer. The possibility that ma
late and oxaloacetate shuttling plays a role in CFM regulation requires fur
ther investigation.