Assimilate partitioning and distribution in fruit crops

In higher plants, leaves function as the principle site of resource acquisition by utilizing the free
energy captured in photosynthesis for the reductive assimilation of oxidized forms of carbon and
nitrogen into carbohydrates and amino acids, respectively. Photosynthate are subsequently
partitioned to the many heterotrophic tissues of the plant with as much as 80 per cent of the
carbon acquired in photosynthesis is transported in the plant’s vascular system to the import
dependent organs. Sucrose is the principal metabolite in this scheme of resource allocation as it
is the major end product of photosynthetic carbon metabolism and, in majority plants; it is the
predominant form of carbon transported to the heterotrophic tissues. This systemic distribution
of photosynthate is known as ‘assimilate partitioning,’ a crucial process associated with plant
growth and productivity.
Source A source of organic material is a region in which organic materials are synthesized.
Example- leaf, root, tuber, or tuber during development.
Sinks Any non-photosynthetic organ or an organ that does not produce enough photosynthate to
meets its own needs. Example- root, seed, fruit, root tuber and tuber during developing.

Leaves are the most important organ for photosynthesis, in which light energy is captured by
green plants (mainly by the chlorophyll in leaves) and used to synthesize reduced carbon
compounds from CO2 and water. Photosynthesis produces carbohydrates for growth and energy
and photosynthates constitute up to 90% of a plant’s dry matter and both growth and cropping
depend on a ready supply of carbohydrates and nutrients.

Source and Sink relationship

The terms source and sink in relation to the transportation of organic molecules in the phloem of
plants.

Food conduction may be in any required direction unlike the water conduction which is
unidirectional process. Pressure flow/ mass flow hypothesis of food/ sucrose translocation –
given by. E Munch (1930). This is the most accepted theory of food conduction in plants.

According to it food translocations occurs in between source and sink in order of turgor pressure
gradient i.e. high turgor pressure to low turgor pressure.
3. Phloem loading/ sucrose loading at source-
It is an active process helped by carrier molecules. At source due to phloem loading
concentration of sieve cells increase, results in increase in osmotic pressure and water will moves
from nearby xylem into sieve cells results in increase in turgor pressure and increase in water
potential. It is establish a higher T.P. at source and sieve tubes. Sucrose moves from source in
sieve tubes towards sink from high T.P/ High water potential to towards the low T.P./Low water
potential.

Phloem unloading/ sucrose unloading at sink

It is an active process helped by carrier molecules. At sink sucrose is unloaded results in
decrease in O.P., it results in exit of water into nearby xylem leads to decrease in T.P. and water
potential of phloem. In sink cells the unloaded sucrose is either changed into starch or consumed,
to maintain low O.P. and contiuous unloading.
So the process of sucrose loading at source and unloading at sink continues. This turgor pressure
difference will maintained and water will continue to move in at source and out at sink.

Passive v. active transport

Passive transport
 Movement down the electrochemical gradient
 From a more positive electrochemical potential
 To a more negative electrochemical potential

Active transport
 Movement against electrochemical gradient
 From a more negative electrochemical potential
 To a more positive electrochemical potential
Path of water absorption
Soil solution→Root hairs→Epidermis→Cortex→ Endodermis→ Pericyclecells→ Protoxylem→
Metaxylem

The path of water from root hair to cortex, may be apoplastic or symplastic. Casparian strips
blocks the apoplast, thus water must passes through passage cells via symplast.
Symplast
A sustainable living path is known as symplast. This is the living passage. The movement of
water from cell to cell through plasmadesmata is called symplastic path in plant.
Apoplast
This is non-living path in plants. Watered cell wall, intercellular space and xylem cavity
bassociate to getherbto form apoplast.

Assimilate partitioning
Assimilate supply is dependent on photosynthesis. The distribution of assimilate determines the
amounts and patterns of plant growth and yield. Translocation dependent on developmental state
of the plant. Transport direction and volume depend on sink position and relative attraction
strength. 90% of sap solute molecules are carbohydrate that travel at a speed of about 50-100
cm/h. Sucrose is main form of translocation. Partitioning within a tree is not a genetically
programmed process, but a result of the unique combination of competing organ and their
relative abilities to compete for limited carbohydrates. The degree of competition among various
sinks depends on the organ activity and distance from the carbohydrate source.
Partitioning priorities and sink competition
Herbaceous crop plants accumulate photosynthates in source leaves during the photoperiod and
evacuate them during night, leaving the leaf "empty" toward morning. Competition for
photosynthate is evident among different organs (e.g., fruit-shoot) as well as among individual
units of the same type of organ (e.g., fruit-fruit). In citrus, the spring flush gives rise to vegetative
shoots, leafy inflorescences, and pure, leafless inflorescences. Shoot elongation and leaf
expansion occur mostly before anthesis and fruit set; direct competition is thus prevented.
Moreover, leafy inflorescences reveal higher rates of fruit set and persistence, indicating that the
leaves support the reproductive organs by provision of photosynthate, hormones, or some other
mechanism. The leaves close to developing fruits exhibit increased photosynthetic capacity as
compared to the remaining leaves of the tree. The calyx has significant photosynthetic capacity,
so in the cape gooseberry, the green calyx that completely covers the fruit during its development
plays an important role in the production and translocation of carbohydrate during the first 10-20
days of fruit development.
On the other hand, in the presence of a heavy crop the vegetative summer flush is poor or absent
altogether, suggesting sink priority of the developing fruit. The retardation of root growth during
periods of shoot flush emergence has been interpreted in terms of root-top competition for
photosynthate, with tops having the priority. Competition between fruit is apparent in citrus, as
in other fruit trees. The progressive reduction in fruit numbers during early fruit development
(fruitlet abscission) has been linked to the carbohydrate status. The inverse relationship between
fruit number and size is another facet of fruit-fruit Competition.
Source-sink manipulations
Fruit load adjustment improves the fruit quality in same year and ensures the accumulation of
reserves which can positively influence tree development for subsequent years. But alternate
fruit bearing is a major problem that can result in serious economic losses for fruit producers. A
high fruit load probably main cause of alternate bearing.
Heavy crop load, as occurs during the "on" year of alternate-bearing cultivars, involves depletion
of both carbon and mineral reserves which may culminate under extreme conditions in tree
collapse. Photosynthate productionis often unable to satisfy the demands during fruit set and fruit
growth following heavy and prolonged.
Fruit removal in apple trees favors more leaf area development compared to those with intact
fruits and subsequent fruiting in young trees reduces leaf area. Furthermore, fruitless growing
strawberries produced 61.1% assimilates in leaves, but only 39.2% and 21.1% of the assimilates
occurred in plants growing with 6 and 12 fruits, respectively. Defoliating trees partially increases
the rate of photosynthesis in the remaining leaves because they provide a relatively larger sink
and this depends on the defoliation degree.
Some special practices which divert the assimilates to developing organs
1. Root prunning- removel of roots 40cm. Away from the plant. Exam. Guava and Citrus
2. Ringing- removel of complete ring of bark from a branch or a trunk. Exa. Mango and grapes
3. Dehorning – to removel of overcrowding & intermingling of branches.
4. Notching – partial ringing of a branches above a dormant lateral bud. Exa. Poona fig
5. Nicking- partial ringing of a branches below a dormant bud. exa. Apple and Poona fig
6. Bending- bending of branches or shoot. Exa. Guava
7. Leaf prunning- removal of old and senescence leaves. Exa. Datepalm.
8. Skirting- removel of low hanging branches. Exa. Mango

Leaf area index (LAI)

LAI in conjunction with sunlight interception is useful as a basis for analyzing canopy
productivity. Apart from cultural practices, agro-ecological conditions and age of plants can
influence LAI development. The rapid LAI development at 2,300 m a.s.l. allowed early and
higher fruit production throughout the culture compared to the higher site. The leaves close to
developing fruits exhibit increased photosynthetic capacity as compared to the remaining leaves
of the tree.

Factor affecting net photosynthesis and photosynthate translocation

Temperature plays an important role in CH partitioning. The optimum temperature for
transporting CH in most species is between between 20 and 30 °C and, and, according to
Guardiola and Garcia-Luis (1993) translocation diminishes with decreasing temperatures (due to
the viscosity of the phloem solution); however, in species not sensitive to low temperature
conditions, the sieve tubes are functional at temperatures close to the freezing point and even
lower. Night temperature is of great importance for carbohydrate translocation. This is because
CH are translocated during night hours and ther fore, as in the case of the Rosaceae, it has been
reported that most growth occures during night than day.

Water stressed plants delay CH transport due to an increase in the viscosity of the solution
translocated. Prolonged water deficits cause the accumulation of abscisic acid, a hormone that
inhibits phloem loading in leaves (Guardiola and García-Luis, 1993). The distribution of
assimilates may be affected by a deficiency or imbalance of mineral nutrients and, furthermore,
by the initiation and development of sink organs and for source functioning, the plant requires an
adequate supply of nutrients. Potassium is claimed to be essential in the process of loading and
unloading the phloem (due to high concentrations of K in companion cells of sieve elements).
Potassium deficiency affects vegetative growth because the plant alters the distribution of K to
improve the growth of the fruit (Teiz and Zeiger 2006)].

Future Prospects
Source sink relationships of plants have become one of the most exciting research areas in recent
years. The subject encompasseas broad array of physiological and biochemical processes, with
significant crop management ramifications. Assimilate partitioning in plants is controlled by a
number of factors that include photosynthesis, the number and location of competing sinks,
storage capacity and vascular transport. Although there is considerable knowledge on individual
processes in plants such as photosynthesis, translocation and cell growth, it appears that the
controls actually regulating the assimilate partitioning at the whole plant level are still poorly
understood (Wardlaw 1990; Le Roux et al. 2001) [32, 33]. Indeed, many processes are closely
interrelated and more integrative research work based on modelling is greatly needed.