15 _
crops such as wheat, barley, oats, and 0-
0
Oa
corn, tomato, and rice have occurred
O c 12 without any increase in relative growth
co._ rate of young plants (14). Over the his- 0
5I
c
potential of field crops, it has been the Q
0-
7 * partitioning of photosynthetic product 0
4
0
.* between economic yield and the rest of C
_
00
0
/ the plant ["harvest index" (HI) (14)] that c
6 0 has been of primary importance, even ._
co CL 3
'C
* though selection was not directed specif-
0
5 0
ically to that end. Harvest index, also 0.5
termed coefficient of economic yield,
.c
b. i 1 4~~~~~~~~~~~
- can apply to the ratio of harvested dry
0.5 [ weight to total aboveground dry weight MG 0.4
(shoot HI) or to the ratio of harvested
0.V
S
/ dry weight to above- plus belowground CD
0
0 0.4
- dry weight (plant HI). In the field, as- I 0.3
0
(D
0 0 sessing root dry weight can be extremely
difficult, so shoot HI is more commonly
0.3 used in agronomic studies. 0.2
1940 1960 1980
1900 1940 1 980 Cultivar introduction date
Cultivar introduction date Harvest Index Fig. 2. Comparative yield potential and HI for
Fig. 1. Comparative yield potential and HI for four major peanut cultivars from the southern
eight British winter wheat cultivars, plotted Improved HI was responsible for the United States, plotted against the year that
against the year that each cultivar was intro- grain yield potential increases among each cultivar was introduced. The cultivars
duced. The cultivars were grown in intensive- successively developed cultivars of the were grown in intensively managed experi-
ly. managed experimental plots at a high- mental plots in a favorable environment near
fertility site near Cambridge, England, in the major cereal species over the past centu- Gainesville, Florida, in the 1976 season.
1977-1978 season (16). ry (1, 14, 15). Eight major winter wheat [Adapted from W. G. Duncan et al. (3)]
802 SCIENCE, VOL. 225
in the field are difficult to obtain, HI -
simply measurable on a crop stand at the
end of the season without destroying the
nwxt generation of seed. Furthermore, in
pijing wheat, HI for spaced plants ap-
p&irs to be representative of the parame-
te fbor a community of plants (18).
Beders require easily measured selec-
tion criteria because of the large popula-
tions to be screened and the relative
difficulty of work in the field; they fre-
quently need to use spaced plants in the
early stages of new cultivar develop- 150
Days after planting
ment.
Recognition of the importance of parti- Fig. 3. Time course of total plant dry weight production ("total") and fruit growth ("pod") for
tioning of fixed carbon between alterna- the peanut cultivars grown in the study represented in Fig. 2. The cultivars are Dixie Runner
tive sinks in past improvements in genet- (released about 1943) and Early Bunch (1977). [Adapted from W. G. Duncan et al. (3)]
ic yield potential has led to much re-
search into the physiology and biochem- remain methodological uncertainties, tion or genetic engineering to suppress
istry involved. Figure 4B shows the main and figures based on other methods oxygenase activity while maintaining or
sinks for the gross quantity of carbon range as high as 50 percent (20). Since enhancing carboxylase activity. This
fixed in the leaves. It illustrates vividly the function of this apparently wasteful would have the double advantage of both
what a small proportion of carbon fixed partitioning is unknown, much research reducing the competitive effect of 02 at
is finally harvested. Figure 4A shows the has been directed at its reduction or the active site and reducing the photores-
typical fate of annual solar radiation inci- elimination by chemical or genetic piratory release of CO2. However, no
dent on crops. Carbon partitioning phe- means. one has been able to refute the argument
nomena start at the very point of carbon The photorespiratory pathway arises that the oxygenase reaction is a physio-
fixation with partitioning between the from an oxygenase activity possessed by chemically inevitable consequence of the
photosynthetic carbon reduction (PCR) ribulosebisphosphate carboxylase (21), carboxylase mechanism (23). Neverthe-
cycle and the photosynthetic carbon oxi- which catalyzes the primary CO2 fixa- less, recombinant DNA methodology
dation (PCO) cycle. They continue to be tion of photosynthesis. The substrate for provides a means for systematic manipu-
expressed in all aspects of carbon econo- both oxygenase and carboxylase activity lation of the amino acid composition of
my while harvestable yield is being gen- is the photosynthate intermediate ribu- ribulosephosphate carboxylase. This, to-
erated. For example, carbon is parti- lose-1,5-bisphosphate. This competition gether with investigation of the exact
tioned between starch storage in the between 02 and CO2 for the active site mechanisms of carboxylation and oxy-
chloroplast (for later mobilization) and in ribulosebisphosphate carboxylase re- genation, offers hope of creating an im-
immediate export into the mesophyll cell duces the gross rate of CO2 fixation (Fig. proved carboxylase/oxygenase ratio for
cytoplasm; between sucrose retention in 4B). A product of the oxygenase reaction crop species or at least of determining
leaf mesophyll and loading into the phlo- is phosphoglycolate. It is the subsequent why it is not possible to do so.
em for export; between retention in the multistep conversion of phosphoglyco-
phloem and unloading into growing late to phosphoglycerate, an intermedi-
sinks; between vegetative growth and ate required in the PCR cycle, that in- Starch Versus Sucrose
reproductive growth; and among com- volves release of photorespiratory CO2. Synthesis in Leaves
peting alternative sinks, such as root and This photorespiratory pathway also ac-
shoot, elongating wheat stem and ear complishes a partial recovery of carbon Early products of the PCR cycle in the
growth, or grains within an ear. Little is partitioned into phosphoglycolate. At- chloroplast are the triose phosphates,
known about the regulation of these pro- tempts to chemically inhibit phosphogly- phosphoglycerate, and dihydroxyace-
cesses. Given the numerous carbon par- colate metabolism to prevent the CO2 tone phosphate. These sugar phosphates
titioning steps that occur between pri- release have not been successful at in- are biochemically and spatially parti-
mary fixation of CO2 and final accumula- creasing net CO2 fixation: usually net tioned between two major biosynthetic
tion of a small fraction of the fixation fixation decreases instead (13). It seems pathways, one leading to the synthesis
product into harvested organs (Fig. 4B), that inhibition of any step in phosphogly- and retention of starch in the chloroplast
only a few aspects can be dealt with colate metabolism prevents a necessary and the other to sucrose synthesis in the
here. recycling of carbon back into the PCR cytoplasm. Up to 50 percent of the pho-
cycle, while the accumul4ted photorespi- tosynthetically fixed carbon can be allo-
ratory intermediates havve no capability cated either to starch or to sucrose de-
PCR Cycle Versus PCO Cycle of feedback regulation of the partitioning pending on several factors, including
between oxygenation and carboxylation plant species, environment, nutritional
Concurrent with photosynthetic car- of ribulosebisphosphate (22). This con- status, and developmental stage of the
bon fixation by the PCR cycle, the inter- clusion focused attention on the ratio of plant (24, 25). This partitioning is impor-
linked PCO cycle releases back to the carboxylase to oxygenase activity of ri- tant to plant growth because the forma-
atmosphere CO2 that was recently fixed. bulosebiphosphate carboxylase. Evi- tion of sucrose (the principal phloem
In C3 species the proportion of CO2 fixed dence suggesting that the ratio is not transport sugar in most crops) is a prime
by the PCR cycle that is partitioned back immutable has encouraged a search in determinant of carbon export from pho-
to CO2 by the PCO cycle is usually,found several laboratories for suitable manipu- tosynthesizing leaves and because leaf
to be 15 to 20 percent (19), but there lations of the protein structure by muta- starch is a major carbohydrate reserve
24 AUGUST 1984 803
that is mobilized to sucrose when current ose phosphates across the chloroplast the cytoplasm during sucrose synthesis
photosynthesis is low relative to sink envelope to the cytoplasm is stoichio- favors continued triose phosphate export
demand for photoassimilate, as occurs in metrically and obligatorily coupled in a from the chloroplast by counterexchange
low light or darkness (26) or, in the one-to-one counterexchange with inor- through the phosphate translocator.
longer term, when the leaves are senes- ganic phosphate (Pi) through this specific Thus, under conditions that favor su-
cing (27). translocator protein (28). In the cyto- crose synthesis, triose phosphates are
Dynamic and balanced partitioning of plasm, sucrose is synthesized from triose partitioned away from the starch biosyn-
the PCR-derived triose phosphates be- phosphates. Key enzymes involved are thetic pathway that resides in the chloro-
tween starch synthesis in the chloroplast fructose-bisphosphatase (FBPase) and plast. If sucrose synthesis in the cyto-
and sucrose synthesis in the cytoplasm sucrose-phosphate synthase (SPS) (29). plasm is reduced, triose phosphates re-
appears to be mediated by the levels of Sucrose synthesis is a phosphate-liberat- main within the chloroplast for starch
certain key metabolites and by the so- ing process (net reaction, 4 triose phos- synthesis. The resulting increase in
called phosphate translocator protein in phates + 3 H20 - 1 sucrose + 4 Pi). phosphoglycerate within the chloroplast
the chloroplast membrane. Export of tri- The liberation of inorganic phosphate in stroma (high phosphoglycerate to Pi ra-
tio) also favors starch synthesis by allo-
sterically activating the starch-synthesiz-
A ing enzyme adenosine diphosphate-glu-
cose pyrophosphorylase (30).
Studies on source-sink manipulations
of whole plants show that photosyntheti-
cally fixed carbon can be preferentially
ents (50 to 56)////f partitioned into sucrose available for ex-
port during periods of high sink demand
or retained by starch when sink demand
is low (31). However, in other plants,
export from photosynthesizing leaves
,oyveswron(60 se
o,//
to 7%~~ continues undiminished in response to a
rapid change in sink demand, at least in
4) the short term; the exported carbon ac-
5%)
cumulates in alternative sinks (32).
Fig. 4. (A) Fate of so- Starch synthesis and accumulation in
Ineffective absorption due to light saturatlion (20 to 40%) lar radiation incident leaves may be controlled indirectly by
over 1 year on a field the rate of sucrose synthesis. A growing
used for a typical body of evidence (29, 33) suggests that
grain crop in a tem- the FBPase and SPS are key regulating
perate agricultural
environment. (B) Par- enzymes in sucrose synthesis. As such,
titioning of gross pho- the activities of these enzymes, when
tosynthetic CO2 fixa- acting in coordination with the phos-
tion by a generalized phate translocator, may represent an im-
temperate grain crop.
B The values in paren- portant link between sink demand and
theses and relative rates of carbon partitioning into starch
width of the bars are and sucrose. For example, the level of
intended as guides to SPS in plants appears to be negatively
'iontopotential)~
X///,,,,,,,,,,,,,,,S,/////
the proportion of the
previous fractional correlated with the total starch content
flow remaining that is of leaves and with biochemical partition-
diverted into each ing of photosynthetic carbon into starch.
>OSe (1,51 to 20%) successive sink. For- Plants that form little starch, such as
aging by insects has wheat, barley, and spinach, have high
not been included be-
//////////////// cause of its extreme SPS activity compared to plants that
0 to 0variability. form large amounts of starch, such as
tobacco, peanuts, and soybeans. Intra-
specific differences in SPS activity and
carbon partitioning have been noted in
wheat (25). Recent studies of soybeans
XI0 toa 20%) 1,17.74, showed that altering the source-to-sink
ratio by partial defoliation, pod removal,
or changes in photosynthetic irradiance
zS/,,,,,,,,,,,,,,,,ble
pa rts (50 to 7 0%b'
. .........X at hoar,ves, ,3Z
caused changes within hours in extract-
~~~~Decayed before harvest M? able SPS activity and in starch and su-
crose synthesis (24, 34). These results
support the hypothesis that SPS activity
is important to starch-sucrose partition-
ing in leaves.
Regulation of FBPase is receiving in-
creased attention with the recent discov-
804 SCIENCE, VOL. 225
ery of fructose-2,6-bisphosphate (FBP) Utilization of the high-energy interme- Vegetative Versus Reproductive or
in plants. FBP plays an important regula- diates from respiration may be opera- Storage Growth
tory role in glycolysis and gluconeogene- tionally divided into that required for the
sis in animal liver (35). In plants the level processing of stored carbohydrate to The observed increase in HI for sever-
of FBP responds to changes in light, new growth and that required for main- al improved crop species reflects a shift
specific metabolites, sugars, and CO2. taining existing cells in a viable state from excessive vegetative development
That FBP is a potent inhibitor of cyto- (41). While it has been argued that the in traditional varieties to greater parti-
plasmic FBPase and sensitizes FBPase biosynthetic pathways that convert glu- tioning into fruits or other sinks for pho-
to the effects of FBP and Pi suggests that cose to the various lipids and nitroge- toassimilate, such as potato tubers or
it plays a key regulatory role in sucrose nous monomers required for growth are swollen storage roots, in modern culti-
biosynthesis (36). efficient (42), energy required for mainte- vars. The basis for this change can be
nance of existing cells may be as much as examined from two perspectives. The
50 percent of the overall respiration rate first concerns the timing and establish-
Catabolism Versus Anabolism in whole plants, and the utilization of this ment of large -numbers of fertile flowers
energy is poorly understood. If the con- or other storage organ initials. The sec-
Energy for growth and maintenance of tinuous breakdown and resynthesis of ond concerns regulation of transport and
plant structure and for ion uptake is existing cellular compounds (turnover) partitioning of current photoassimilate
derived largely from respiratory catabo- could be slowed, or if the energetic cost and prior reserves into these developing
lism ofphotosynthetically derived sugars of maintaining inter- and intracellular harvestable organs rather than into fur-
to CO2 and water (37). Up to 50 percent chemical and electrochemical gradients ther branching, rooting, or ineffective
of the net CO2 fixation by leaves of an across membranes could be reduced, flowering.
annual plant may ultimately be lost to the more carbon might be available for eco- For traditional varieties of species
plant by subsequent respiration. Crop nomic yield. from which seeds are harvested, such as
scientists have long questioned whether While improvement of respiratory effi- cereals and grain legumes, the number of
the partitioning of photosynthetically ciency may not have the potential for flowers formed and fertilized far exceeds
fixed carbon between biosynthesis very large yield increases, there is evi- the number that fill to mature seeds.
(anabolism) and respiration (catabolism) dence of increased plant growth rate and Despite having less reproductive redun-
could be shifted in favor of more biosyn- yield in association with decreased respi- dancy, modern high-yielding cultivars
thesis. Can respiratory efficiency be im- ration. In corn and tall fescue genotypes still have much fruitless flowering. In
proved, and if so, would it improve crop possessing high growth rates, low rates terms of yield components, however, the
yield? of leaf respiration were found (43). More improvement during the era of systemat-
Two possible routes to greater effi- directly, selection for intravarietal varia- ic breeding has mostly involved number
ciency in respiratory utilization of fixed tion in respiration rate of mature leaf of fruits harvested per hectare rather
carbon can be envisioned. Either the segments of perennial ryegrass readily than fruit size.
efficiency of conversion of stored carbo- gave lines with high or low respiration A key to high yield is the establish-
hydrate to adenosine triphosphate might rates. Measurement of forage yield under ment of a large number of seeds before
be increased or utilization of adenosine simulated cropping conditions in both their rapid filling commences. Barring
triphosphate for processes not strictly greenhouse and field studies (44) showed extraordinary weather or other calamity,
required for plant growth in the crop that the lines with low respiration rates once seed number has been established
environment might be reduced. had a consistent 6 to 13 percent greater mature yield has largely been predeter-
While bioenergeticists traditionally annual dry matter yield than the parent mined. The average mature weight per
have viewed respiration as an optimized population from which they were de- seed is a more conservative property of a
process under tight control, in plants rived. This increase came primarily from cultivar than is seed number per hectare.
there is an alternative terminal oxidase later summer cuttings in a continuous These generalizations can be illustrated
that transfers electrons from the cyto- cropping regime, when growth tempera- with wheat. Two weeks before pollina-
chrome oxidase chain to oxygen at ubi- ture was relatively high and the amount tion starts, a modern dwarf wheat has
quinone. This cyanide-insensitive alter- of root and stem tissue was large. As several times more partly developed
native oxidase is not coupled to adeno- such, it may indicate that some portion flowers than will set and fill grain (46).
sine triphosphate synthesis (38) and as of the maintenance respiration require- Each spikelet (that is, inflorescence
such appears to be wasteful of energy. ment was decreased in these lines. branchlet) in the central part of the spike
The pathway is found in many species, in The metabolic basis for this reduction can have nine to ten primordial florets.
both shoots and roots (39, 40), but does in respiration is not known, nor is there However, about two of these florets
not operate until the capacity of the any indication that yield increases asso- have ill-formed anthers'that render them
cytochrome oxidase system has been ciated with it would necessarily follow if incompetent to set grain. During the 2
exceeded or blocked after ubiquinone. genotypes with such reduced respiration weeks before pollination there is a col-
Considering the degree of engagement of were established in annual grain crops. lapse in floret competency, such that
the pathway in roots, it was calculated About half of the yield increase was typically only two or three florets will
that in wheat the carbon loss to that associated with an improved ability of actually set grain. This 60 to 70 percent
pathway was at least 6 percent of the the lines with low respiration rates to loss of competency to set grain appears
carbon in final grain yield (39). If a loss establish new leaf cover after defoliation to be closely related to the supply of
of such magnitude occurs and can be (45). So the yield enhancement from a photosynthetic assimilates at the time.
shown not to be associated with an es- reduction in respiration rate might not be Consequently, in the 10- to 20-day period
sential function, its elimination could significant in annual grain crops, which before pollination final seed number is
translate into a significant increase in the are not required to undergo multiple re- particularly sensitive to irradiance. Low
partitioning of carbon to economic yield. growth. light at that time can cause an irretriev-
24 AUGUST 1984 805
able loss of yield because fewer seeds mesophyll cells; (iii) intracellular com- occur at the carrier level or by a cell
are established (47). As a consequence of partmentation between cytoplasm and turgor-dependent process in which in-
this sensitivity of floret abortion to pho- vacuole; (iv) compartmentation between formation about the rate of unloading in
toassimilate supply in temperate cereals, the symplast (or intracellular volume) sinks is relayed instantly to the source as
CO2 enrichment to boost photosynthesis and apoplast (or extracellular "free a hydrostatic pressure change in the
of field crops before anthesis can raise space" in cell walls and intracellular whole transport network or by changes
final yield by increasing grain number spaces); and (v) accumulation of solutes in the rate of water entry into the sieve
per unit of ground area (48). By contrast, by the sieve element-companion cell elements (54).
CO2 enrichment after anthesis has a less- complex of the vascular tissue. The size, The sieve elements form a continuous
er effect on yield. distribution, and composition of these transport network throughout the plant,
Although the main component of vari- various pools varies with species and with numerous branches and anastomo-
ation in yield of wheat (and other spe- with translocation status of the plant. In ses. There is no evidence of one-way
cies) is the seed number per unit of soybean leaves, for example, there is a valves, so in principle any source leaf
ground area, mean seed weight can nev- preferential net accumulation of starch in can supply solutes to any sink without
ertheless vary somewhat by improved the second palisade mesophyll layer dur- leaving the sieve tube network. Experi-
photoassimilate supply (48, 49) and cer- ing the stage from flowering to early seed ments with radioactively labeled solutes
tainly differs among genotypes. Just as filling. This starch does not turn over show that in some species there are
kernel number is established before rap- diurnally. It is a longer term reserve, preferred routes from certain source
id grain filling gets under way, so is the being mobilized during the middle to late leaves to certain sinks but that selective
average potential size of the seed. This is stages of seed filling, when leaf photo- leaf or sink removal rapidly alters the
closely related to the mean number of synthesis has markedly declined (27). pattern of movement in most species (15,
cells per seed in other species as well as A portion of the sucrose synthesized 55). It seems unlikely that source leaves
cereals (50). Cell division in the starchy through FBPase and SPS in the cyto- have mechanisms to determine where
endosperm of wheat is virtually com- plasm is stored in the vacuoles. Kinetic their solutes are destined (56). General-
plete within 2 weeks after pollination studies (53) show that the cytoplasmic ly, the preferred channels of transport
(51). Once kernel number per unit of (or transport) sucrose pool has a much seem to be associated more with the
ground area and cell number per kernel faster turnover than the larger vacuolar least resistive vascular route between
are established, the filling of the grain pool, which is less available for immedi- source and sink (15).
usually proceeds rapidly (in about 4 to 6 ate export through the mesophyll toward Although the mechanism of long-dis-
weeks) to a substantially predetermined the veins. The vacuolar sucrose pool, tance transport has not been unequivo-
mean kernel size. From that perspective, exchangeable with the cytoplasmic pool, cally proven, the consensus is that there
by 2 weeks after grain set the partitioning is the first source for export during the is a bulk flow of solution along sieve
of photoassimilate appears to be largely night. Starch appears to be mobilized to tubes under the influence of a hydrostat-
regulated by the filling sinks themselves. sucrose at night only when the vacuole ic pressure gradient. This gradient is
The physiology behind such regulation pool is substantially depleted. maintained osmotically by regulated
has attracted much research: regulation The sieve element-companion cell loading of solutes into the sieve element-
of CO2 fixation rate per unit of leaf area complex of the phloem contains a much companion cell symplast in source re-
by sink demand; control over the or- higher concentration of sucrose than the gions and regulated unloading in sink
dered senescence of the leaf canopy by surrounding tissues. The partitioning regions. Measurement of sieve element
storage sinks; and regulation of the day- step leading to that high concentration is sucrose concentrations and gradients in
to-day partitioning of current photoassi- called phloem loading. The following sugarbeets, soybeans, and other species
milate between competing alternative sketch, based on detailed reviews (54), is showed them to be sufficient to generate
sinks. The last of these topics has been our currently favored view of how phlo- the necessary hydrostatic pressure gradi-
studied phenomonologically by estab- em loading occurs. ent (57). Long-distance transport seems
lishing principles of competition between The intricately reticulated network of unlikely to impose appreciable limitation
sinks of different sizes and distances veins in leaves ensures that no more than on the rate of transport from sources to
from the source (52). It is also being two or three cells need be traversed sinks even when partial incisions or re-
studied by tracing the mechanisms and between photosynthetic mesophyll cells strictions are made in the dominant
potential points of regulation of carbon and sugar-accumulating phloem cells. transport routes (15, 58)-alternative
compound flow between chloroplasts in Photoassimilates, mostly sucrose, travel paths appear to take over.
the leaves to uptake in cells of the devel- symplastically down their concentration The discussion so far suggests that
oping sink. gradients to the mesophyll cells close to photosynthesis and the mechanism of
the companion or phloem parenchyma phloem loading determines the amount
cells, where they are released without of photosynthetic assimilate made avail-
Carbon Transport from Source to Sink hydrolysis by a facilitated efflux mecha- able for translocation, whereas the
nism into an apoplastic solution of rela- mechanism and kinetics of unloading
Complementary to biochemical parti- tively low concentration. An energy-re- into, and associated uptake by, compet-
tioning between sucrose and starch is quiring active process then loads sucrose ing sinks, in association with relative
physical compartmentation of these into the phloem, creating a high sucrose distances between sources and sinks,
products in the leaf. Histochemical and concentration in the sieve tube symplast. determines the partitioning of loaded ma-
kinetic studies show at least five major This energetically "uphill" loading of terial in the short term.
types of photoassimilate compartmenta- sucrose appears to occur, at least in part, Less is known of the regulation and
tion in leaves: (i) differential starch stor- by a plasmalemma-bound, sucrose-spe- coordination of phloem unloading (solute
age among the different leaf cell types; cific carrier involving a sucrose-proton efflux from the phloem and interconnect-
(ii) different sucrose storage pools among cotransport mechanism. Regulation may ed tissues of the vascular bundles) than
806 SCIENCE, VOL. 225
of the subsequent uptake and assimila- mercially useful sinks. In grain crops developments through genetic engineer-
tion of solutes for growth and storage improving crop photosynthesis rate by ing, and although yield-enhancing chemi-
product formation in sink tissues. The environmental amelioration is most cals hold promise, the complexity of
diversity of sink types suggests that effective at increasing yield (by increas- yield and our lack of understanding of
there are several mechanisms of unload- ing the number of grains competent to plant physiology and development are
ing. fill) if effected before grain filling starts. such that the rational design of such
Unloading from the vascular bundles Increasing sink size (grain number per approaches for yield improvement is
can be through symplastic plasmodesma- hectare) by chemical or breeding meth- only in its infancy.
tal connections to growing cells, as in ods is likely to be an effective route to
pea and corn roots (59) and in young higher yields of seed crops; this will References and Notes
leaves (60), or unloading may be across a involve prevention of abscission or abor- 1. K. J. Frey, in Genetic Engineering for Crop
Improvement, K. 0. Rachie and J. M. Lyman,
cell membrane into the apoplast of the tion of flowers and young seeds while Eds. (Rockefeller Foundation, New York,
sink cells, as in sugarbeet tap roots (61) retaining the ability to fill these extra 1981), p. 15.
2. D. N. Duvick, Maydica 22, 187 (1977).
and in the storage stem of sugarcane sinks. 3. W. G. Duncan, D. E. McCloud, R. L. McGraw,
(62). In crop seeds solutes may first pass The rate of growth of established, de- K. J. Boote, Crop Sci. 18, 1015 (1978).
4. S. H. Wittwer, Science 188, 579 (1975).
symplastically from the phloem through veloping seeds seems to be determined 5. J. L. Monteith, Philos. Trans. R. Soc. London
Ser. B 281, 277 (1977).
one or more maternal seed coat tissues primarily by phenomena operating either 6. A. F. Hawkins, Outlook Agric. 11, 104 (1982).
before entering the apoplast of the em- at the stages of unloading of photoassi- 7. T. J. Johnston, J. W. Pendleton, D. B. Peters,
D. R. Hicks, Crop Sci. 9, 577 (1969); J. B.
bryonic sink. Furthermore, solutes (prin- milate into the extracellular solution sur- Schoper, R. R. Johnson, R. J. Lambert, ibid. 22,
cipally sucrose and amino acids) may rounding seed cells or accumulation and 1184 (1982).
8. One hundred percent light interception is im-
remain unaltered during unloading or, conversion to storage product of this practical. The leaf area index giving 95 percent
alternatively, be partially metabolized unloaded photoassimilate. However, if interception is termed critical leaf area index,
being that which for practical purposes gives
after (or during) unloading (63). HI is further increased, the ability of the maximum growth rate and is taken to constitute
full interception [R. W. Broughham, Aust. J.
The seed coat of the developing le- reduced photosynthetic surface per sink Agric. Res. 7, 377 (1956)].
gume seed is proving to be a convenient organ to supply the necessary photoassi- 9. D. B. Egli, J. W. Pendleton, D. B. Peters,
Agron. J. 62, 411 (1970); R. M. Gifford, in
system for studying unloading (64). In milate will be reduced, leading to a great- Photosynthesis, vol. 2, Carbon Metabolism and
that system, the phloem reticulates to er prospect for improvement in net pho- Plant Productivity, Govindjee, Ed. (Academic
Press, New York, 1983), p. 459.
varying degrees, depending on species, tosynthesis rate per unit of leaf area 10. This may not be true when attempts to increase
light interception by high-density planting are
throughout the seed coat, supplying as- being expressed in higher grain yield. made. For example, maize can suffer reduced
similates to the embryo across the apo- In leaves, partitioning of fixed carbon grain yield due to infertility in high-density
plantings; sugarbeets may have a reduced sugar
plast separating the two generations. Un- between its retention in the plant and its yield in plantings giving a maximum leaf area
loading from the seed coat appears to photorespiratory release is readily main- index greater than 3, even though total crop dry
weight may respond to a leaf area index of S [P.
involve an energy-dependent, carrier- tained by environmental change, but has J. Goodman, Agric. Prog. 20, 1 (1966)].
mediated process (65), rather than just so far defied attempts at genetic alter- 11. A. L. Christy and C. A. Porter, in Photosynthe-
sis, vol. 2, Carbon Metabolism and Plant Pro-
the passive leakage postulated earlier ation. The potential agronomic rewards ductivity, Govindjee, Ed. (Academic Press,
(15), despite a large downhill concentra- for success are so high, however, that New York, 1983), p. 499.
12. J. N. Gallagher and P. V. Biscoe, J. Agric. Sci.
tion gradient from sieve tube symplast to the remaining avenues in that endeavor 91, 47 (1978).
13. R. M. Gifford and C. L. D. Jenkins, in Photo-
embryo apoplast. This suggests that un- must be followed. Conversely, the ap- synthesis, vol. 2, Carbon Metabolism and Plant
loading may serve as a potential control parent scope for improving partitioning Productivity, Govindjee, Ed. (Academic Press,
New York, 1983), p. 419.
point in source-to-sink transfer of photo- of fixed carbon between carbon skele- 14. C. M. Donald and J. Hamblin, Adv. Agron. 28,
synthate and in intraplant competition tons for plant synthesis on the one hand, 361 (1976); L. T. Evans, in Genetic Engineering
of Plants, T. Kosuge et al., Eds. (Plenum, New
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some detrimental consequences follow evidence of an inadvertent increase in photosyn-
thesis energy fixation with successive cultivars.
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to increased commercial yield can be with annual crop species. dry weight (high lipid and protein) is higher than
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