Seasonal Growth Cycle Research Articles

On the importance of sediment-water exchange processes of ammonia to primary production in shallow areas of the Sado estuary (Portugal)

Coupling between the seasonal phytoplankton growth cycle and resuspension mechanisms in low depth areas was studied. Tidal flood in an estuarine inlet with a large area of intertidal flats was investigated with respect to nutrients and suspended particulate matter, during two different periods of the year. A prespring bloom period in March and another during the productive summer term in August were picked. Results indicate that tidal resuspension mechanisms are important in mobilizing NH4+ from the sediment surface pool, and that this additional input of the limiting nutrient is rapidly consumed in the flooding water column, in benefit of microphytes resident in the nutrient-poor, incoming,coastal waters.

Establishment, Spread and Initial Impact of Aphis chloris Koch (Hemiptera: Aphididae), Introduced into Australia for the Biological Control of St John's Wort

The aphid, Aphis chloris Koch, quickly established following field release at numerous sites throughout south-eastern Australia. Local populations built up to levels at which migratory alates were produced within the first season. Dispersal of these alates over large distances led to widespread colonization of St John's wort infested areas within 2-3 years. Detailed observations at the primary release site at Pierce's Creek, ACT, showed that aphid populations undergo a seasonal cycle of population growth throughout the summer, followed by a migratory phase with alate production and dispersal in autumn and a population decline over winter. This pattern was repeated at other release sites monitored over a 5-year period from 1987 to 1991 inclusive, without populations building up to damaging levels. Natural enemies, competition from existing biological control agents, climatic factors and emigration appear sufficient to limit such outbreaks. While caged plants did show a decline in vigour and seed production under heavy aphid attack, such damage was intermittent in open field situations. The present study suggests that, while A. chloris is now widespread, it will not make an important contribution to the overall control of St John's wort.

Meiobenthic Communities Associated with the Seasonal Cycle of Growth and Decay of Ulva rigidaAgardh in the Palude Della Rosa, Lagoon of Venice

Eutrophication in the Palude della Rosa, in the Lagoon of Venice, results from agricultural runoff giving rise to a seasonal cycle of proliferation and subsequent decomposition of the green alga Ulva rigidaduring the summer months, with a dramatic fall in oxygen levels. A survey of the meiofauna at 42 stations in the Palude in December 1991 showed that there were two distinct associations of nematodes and total meiofauna (nematodes plus copepods), one associated with that part of the Palude where Ulvaattains a high biomass, and another where Ulvabiomass is low or absent. Species characteristic of the Ulvaregion include nematodes of the genus Diplolaimellaand copepods of the genus Tisbe, both of which are known from elsewhere to be associated with decaying plant material and implicated in the decomposition process. In the Ulvaregion both diversity (as evidenced from k-dominance curves) and species composition (using non-metric MDS) varied considerably from season to season, associated with the seasonal cycle of growth and decay of Ulva, but diversity and species composition were more stable seasonally in the non- Ulvaregion. For the copepods alone, differences between the Ulvaand non- Ulvaregions were not as evident and species composition in both regions changes seasonally in the same way, although diversity was only reduced markedly in the summer at the Ulvastations. We conclude that it is not only likely that the Ulvacycle controls seasonal changes in meiobenthic community composition, but in turn the meiobenthic community has some control over the Ulvacycle.

Effect of nutrition on seasonal patterns of LH, FSH and testosterone concentration, testicular mass, sebaceous gland volume and odour in Australian cashmere goats

The effects of season and diet on LH, FSH and testosterone concentrations, testicular mass, sebaceous gland volume and male odour were examined in mature Australian cashmere goat bucks fed ad libitum with diets of low or high quality for 16 months under natural photoperiod at 29 degrees S, 153 degrees E (n = 6 per treatment). Each week plasma was sampled, the bucks were weighed, scored for male odour and assessed for testicular mass based on scrotal circumference. Each month a skin sample was taken from the occipital region for histological assessment of sebaceous gland volume. For each variable there was a clear circannual cycle that was significantly influenced by dietary treatment. In bucks fed the low-quality diet, the timing of seasonal changes in LH and testosterone concentration, sebaceous gland volume and odour score was similar, with a mid-autumn peak. In each case the high-quality diet advanced, extended the duration and increased the magnitude of the seasonal increase. FSH concentrations peaked in late spring (in bucks on the high-quality diet) or summer (in bucks on the low-quality diet), reaching a nadir in early winter. The high-quality diet significantly increased concentrations only in the last 2 months of the experiment (spring). There was no overall association between these variables and change in testicular mass; instead, it was strongly correlated with voluntary feed intake and change in body mass, themselves subject to seasonal variation with a winter or spring peak. The high-quality diet induced large increases in body mass and testicular mass during the first months of the experiment without influencing the seasonally low concentrations of FSH, LH and testosterone present at the time. These results demonstrate that the male, like the female, Australian cashmere goat, exhibits marked reproductive seasonality, and that nutrition is a powerful modulator of the seasonal cycle. They suggest that testosterone concentration, sebaceous gland volume and odour score are ultimately dependent upon LH secretion, which appears to be under strong seasonal (photoperiodic) control, with the effects of enhanced nutrition limited to periods when photoperiodic inhibition is waning. However, seasonal regulation of testicular mass, and therefore sperm production, appears to be primarily dependent on changes in voluntary feed intake and growth, with the seasonal cycle of testicular mass more a consequence of the seasonal appetite or growth cycle than of changing gonadotrophin concentrations.

Leaf and canopy photosynthetic CO2 uptake of a stand of Echinochloa polystachya on the Central Amazon floodplain : Are the high potential rates associated with the C4 syndrome realized under the near-optimal conditions provided by this exceptional natural habitat?

The C4 grass Echinochloa polystachya, which forms dense and extensive monotypic stands on the Varzea floodplains of the Amazon region, provides the most productive natural higher plant communities known. The seasonal cycle of growth of this plant is closely linked to the annual rise and fall of water level over the floodplain surface. Diurnal cycles of leaf photosynthesis and transpiration were measured at monthly intervals, in parallel with measurements of leaf area index, canopy light interception and biomass. By artificial manipulation of the light flux incident on leaves in the field light-response curves of photosynthesis at the top and near to the base of the canopy were generated. Fitted light-response curves of CO2 uptake were combined with information of leaf area index, incident light and light penetration of the canopy to estimate canopy rates of photosynthesis. Throughout the period in which the floodplains were submerged photosynthetic rates of CO2 uptake (A) for the emergent leaves were high with a mean of c. 30 μmol m-2 s-1 at mid-day and occasional values of 40 μmol m-2 s-1. During the brief dry phase, when the floodplain surface is uncovered, there was a significant depression of A, with mid-day mean values of c. 17 μmol m-2 s-1. This corresponded with a c. 50% decrease in stomatal conductance, and a c. 35% depression in the ratio of the leaf inter-cellular to external CO2 concentration (c i/c a). During the dry phase, a midday depression of rates of CO2 assimilation was observed. The lowest leaf area index (F) was c. 2 in November-December, when the flood plain was dry, and again in May, when the rising floodwaters were submerging leaves faster than they were replaced. The maximum F of c. 5 was in August when the floodwaters were receding rapidly. Canopy light interception efficiency varied from 0.90 to 0.98. Calculated rates of canopy photosynthesis exceeded 18 mol C m-2 mo-1 throughout the period of flooding, with a peak of 37 mol C m-2 mo-1 in August, but declined to 13 mol C m-2 mo-1 in November during the dry phase. Estimated uptake of carbon by the canopy from the atmosphere, over 12 months, was 3.57 kg C m-2. This was insufficient to account for the 3.99 kg C m-2 of net primary production, measured simultaneously by destructive harvesting. It is postulated that this discrepancy might be accounted for by internal diffusion of CO2 from the CO2-rich waters and sediments via the roots and stems to the sites of assimilation in the leaves.

Age Estimation and Longevity in Reptiles

After a brief recall of the classical meaning of the concept of longevity, the first part of this chapter describes and summarises the main current technique for the estimation of age in reptiles. Among them, sclerochronology is primarily taken into account. The cautious analysis of seasonal growth cycles recorded in hard tissues, although not as rigorous as the mark-release-recapture method of animals in their natural conditions, now appears as a rapid and reliable chronological tool already successfully used in individuals of many reptile species. Sclerochronology is especially efficient for the comparison of several populations, and it is the only method for fossils. The second part presents a synthetic review of known longevities and records in the different groups of reptiles. A short discussion about the significance of longevity shows that for reptiles, because of their thermic metabolism (ectothermy), the physiological longevity must be strongly distinguished from the chronological longevity, especially for a comparison with that known for birds and most mammals.

Mineral uptake and transport in xylem and phloem of the proteaceous tree, Banksia prionotes

Xylem sap of sinker (tap) root, cluster feeding roots, lateral roots and from an age series of main stem extensions of 6-year trees of Banksia prionotes was collected and analyzed for principal organic and inorganic solutes. During the phase of root uptake activity in winter and spring, cluster roots were principal xylem donors of malate, phosphate, chloride, sodium, potassium and amino acid N whereas other parts of the root served as major sources to the shoot of other cations, nitrate and sulphate. Sinker root xylem sap was at all times less concentrated in solutes than that of lateral roots into which cluster roots were voiding exported solutes. Phosphate was abstracted from xylem by stem tissue during winter and it and a range of other solutes released back to xylem immediately prior to extension growth of the shoot in summer. Phloem sap collected from mid regions of stems was unusually low in potassium and phosphate relative to chloride and sulphate in comparison with phloem sap of other species, and its low potassium:sodium ratio relative to xylem indicated poor discrimination against sodium during phloem loading. Data are discussed in relation to the asynchronous seasonal cycles of nutrient uptake and shoot growth.

The influence of birth date on the development of seasonal cycles in red deer hinds (Cervus elaphus)

The relative importance of age and photoperiod in the timing of seasonal cycles was studied in red deer during the first 18 months of life. Seasonal cycles of appetite, live‐weight change, coat growth, oestrus and metabolic rate were examined in two groups of seven hind calves born in early May (group E) or nine weeks later in July (group L).There was no difference between groups in the timing or amplitude of the seasonal voluntary food intake (VFI) cycle, or in the timing of seasonal changes in rate of live‐weight gain, although mean live‐weight gain was significantly greater for group L than group E during the first winter of life.Four animals in group L retained their calf coats throughout the first winter. In group L, the first summer coat fibres appeared approximately four weeks earlier, and mean fibre length for the subsequent winter coat was longer in group L than in group E. There was no difference between groups in the timing or amplitude of the seasonal cycle of plasma prolactin concentration, or in the incidence and mean time of onset of puberty.There was no difference between groups in mean energy expenditure per unit of live weight at any time of the year. Mean daily energy expenditure was significantly higher in summer than in winter, but values measured when the animals were resting and corrected for differences in VFI were similar throughout the year.It is concluded that the seasonal cycles of appetite, live‐weight change and oestrus in young red deer hinds are not modified by age, although the normal cycle of coat replacement can be disturbed in animals born towards the end of the calving season.

Phenotypic Variability in Life History Traits and Phenology of Field Populations of Cardamine flexuosa and C. fallax (Cruciferae) in Honshu, Japan

Abstract Variability in life history traits and phenology of field populations of Cardamine flexuosa and C. fallax was investigated. C. flexuosa is an annual that typically grows all year around and occurs as a weed in cultivated fields such as paddy fields, crop gardens and orchards. C. fallax is a typical winter‐green annual that mainly grows along the roadside on foothills. Differences in growth form of both species corresponded to variations in habitat conditions. Plants of paddy field populations of C. flexuosa normally possess many basal branches, while those of orchard populations of C. flexuosa typically have an erect growth form with few basal branches. C. fallax also possesses a typical erect growth form. Other life history traits (e.g., individual plant biomass and reproductive traits) of both species were highly variable both within and between populations. In C. flexuosa, seasonal differences were also found in flowering and fruiting performance. Synchronous flowering of all plants was observed in spring populations, while partial flowering occurred in summer‐autumn populations. Differences in the size class structure of plants in the autumn appeared to be caused by different cultivation schedules for crop plants in the fields. C. fallax, in contrast, possesses a rather fixed seasonal growth cycle as a typical winter‐green annual with a synchronized flowering in spring.

Prolonged exposure to melatonin leads to time-dependent sensitization of adenylate cyclase and down-regulates melatonin receptors in pars tuberalis cells from ovine pituitary.

In photoperiodic mammals, seasonal cycles of growth and reproduction are cued by changes in the duration of the nocturnal profile of secretion of the pineal hormone melatonin. To investigate the likely mode of action of this hormone on target tissues, the effect of prolonged treatment with melatonin on the sensitivity of the adenylate cyclase (AC) system was examined in primary cultures of ovine pars tuberalis (PT) cells. When cells were exposed to melatonin (100 pM or 1 microM) for 16 h, and the hormone was then removed by a series of washes, basal production of cAMP was elevated over that observed in cells not treated with melatonin. Moreover, the rate of accumulation of cAMP after stimulation with forskolin (1 microM) was markedly enhanced in cells previously treated with melatonin compared to that in untreated controls. This sensitization by melatonin of the basal and forskolin-stimulated responses developed gradually and was half-maximal after approximately 8 h of exposure. There was no significant difference between the sensitizing effects of the two melatonin concentrations used. Treatment with melatonin for 24 h reduced the total amount of specific [125I]iodomelatonin binding in PT cell membranes by 30-50%. However, over the same period there was no reduction in the ability of a maximal (1 microM) concentration of melatonin to inhibit forskolin-stimulated cAMP production, indicating the presence of an excess capacity of melatonin receptors in cultured PT cells. Nevertheless, treatment with melatonin for 16 h did result in a 10-fold increase in the IC50 for the inhibition by melatonin of forskolin-stimulated cAMP production. The enhancement of cAMP production after prolonged treatment with melatonin was not masked by the inclusion of isobutylmethylxanthine (1 mM) during the subsequent challenge with forskolin, suggesting that sensitization was not due to a reduction in the activity of cAMP-phosphodiesterase. In control cells, aluminium fluoride caused an inhibition of forskolin-stimulated cAMP production. Prolonged treatment with melatonin abolished the inhibitory effect of aluminium fluoride, suggesting that treatment with melatonin caused a shift in the net balance between the G-protein-mediated stimulatory and inhibitory influences on the AC system. The sensitization of AC was not blocked by the inclusion of cycloheximide (10 micrograms/ml) during prolonged exposure to melatonin, suggesting that de novo protein synthesis is not a requirement for this effect of the hormone. These results constitute the first demonstration of an independent action of melatonin on ovine PT cells that is dependent upon the duration of the endocrine stimulus.(ABSTRACT TRUNCATED AT 400 WORDS)

Adaptive responses of roots and root systems to seasonal changes

The developmental rhythm and morphology of some perennial herbs in Heilongjiang province, China, have been observed over several years. In the present paper, the development of root systems of mature plants of four species is reviewed. The seasonal growth cycle of root and root system throughout one year is differentiated into three types: (i) renewal type, including partial renewal ( Panax ginseng) and total renewal ( Fritillaria ussuriensis), (ii) continuous growth ( Asarum heterotropoides), and (iii) an intermediate type ( Adonis amurensis). The developmental rhythm of the root system of P. ginseng is co-ordinated with the modern local climate, whereas the developmental rhythms of the other three species are not. These observations indicate that the seasonal growth cycle of a root system can be either the result of an adaptive response of the species to the particular environment in which it grows, or it is the outcome of its past ecological and evolutionary history. Chinese medicinal compounds of economic importance (e.g. ginsenoside) are found in roots and also show seasonal fluctuations.

Seasonal cycles of growth, development and movement of the African honey bee,Apis mellifera scutettata, in Africa

The relationship between the annual colony cycle and seasonal patterns of forage availability was investigated for the African honey bee,Apis mellifera scutellata, in the Okavango River Delta, Botswana. The annual cycle occurred in three distinct periods. The swarming season occurred from October-November, following two to three months of intense brood production, and coincided with the end of peak forage abundance. The migration season occurred from November-May and coincided with reduced and variable floral resources. During the migration season, brood production and food storage were generally low but quite variable from month to month, and swarms passing over the study area at this time traveled in an easterly direction. The migration season was followed by the establishment period (June-September), in which large numbers of new colonies traveling from the west moved into the study area. The establishment period coincided with, and slightly preceded, the period of peak forage abundance, and colonies devoted resources collected at this time almost entirely to brood rearing, which culminated in swarm production. The data suggest that honey bee colonies in the Okavango are mobile and gear their reproduction and movement to seasonally shifting resource pattern.

Endogenous levels of oxygen, carbon dioxide and ethylene in stems of Norway spruce trees during one growing season

The trees sampled in this study came from two stands of Norway spruce, Picea abies (L.) Karst., near Stockholm, Sweden, differing in mean age and height. Holes were bored perpendicular to the stem surface, and gas samples were taken from the outer part of the sapwood throughout one growing season. Endogenous levels of molecular oxygen (O2), carbon dioxide (CO2) and ethylene in the outer sapwood were determined by combined gas chromatography — mass spectrometry (GC-MS) and GC. O2 concentrations began to decrease as growth started in spring. The lowest levels (<5%) were recorded around mid-summer. In the younger stand concentrations remained below 5% until September. In October, O2 concentrations in the sapwood were similar to those of air. Concentrations of CO2 were below 1% in spring, but began to rise in May, peaking a month later at approximately 10%. Thereafter a slow decrease occurred until October, by which time levels had returned to those recorded in spring. Ethylene concentrations in the older stand reached 75 ppm early in May, while levels in the younger stand peaked at around 30 ppm later in May. Thereafter ethylene levels in both stands started to decrease down to ppb levels. The correlation between determined gas levels and physiological events associated with the seasonal growth cycle in temperate zones is discussed.

Survival strategies in arctic ungulates

Arctic ungulates usually neither freeze nor starve to death despite the rigours of winter. Physiological adaptations enable them to survive and reproduce despite long periods of intense cold and potential undernutrition. Heat conservation is achieved by excellent insulation combined with nasal heat exchange. Seasonal variation in fasting metabolic rate has been reported in several temperate and sub-arctic species of ungulates and seems to occur in muskoxen. Surprisingly, there is no evidence for this in reindeer. Both reindeer and caribou normally maintain low levels of locomotor activity in winter. Light foot loads are important for reducing energy expenditure while walking over snow. The significance and control of selective cooling of the brain during hard exercise (e.g. escape from predators) is discussed. Like other cervids, reindeer and caribou display a pronounced seasonal cycle of appetite and growth which seems to have an intrinsic basis. This has two consequences. First, the animals evidently survive perfectly well despite enduring negative energy balance for long periods. Second, loss of weight in winter is not necessarily evidence of undernutrition. The main role of fat reserves, especially in males, may be to enhance reproductive success. The principal role of fat reserves in winter appears to be to provide a supplement to, rather than a substitute for, poor quality winter forage. Fat also provides an insurance against death during periods of acute starvation.

Seasonal Cycles of Growth and Tetraspore Formation in Glossophora kunthii (Phaeophyta, Dictyotales) from Pacific South America: Field and Laboratory Studies

The aim of the present study was: to establish the seasonal patterns of growth and fertility of plants in natural field conditions; to determine whether fertility can be induced throughout the year under constant culture conditions; and to relate these findings to the available data on seasonal variations of environmental parameters

Influence of short-term atmospheric CO2 enrichment on growth, allocation patterns, and biochemistry of black spruce seedlings at different stages of development

Black spruce seedlings (Piceamariana Mill.) were exposed to either elevated (1000 ppm) or ambient (340 ppm) atmospheric CO2 levels at different stages of seedling development over a winter greenhouse production cycle. Seedlings germinated in early February and were placed in CO2 chambers for either 3 or 6 weeks during March, April, May, or August. Total seedling biomass increased under high CO2 conditions for the March, April, and May stages of development, but showed no significant response in August. The greater part of the CO2 response occurred during the second 3 weeks of exposure in March and April but during the first 3 weeks of exposure in May. In September, those seedlings exposed to CO2 in April and May had 30 and 14%, respectively, greater biomass than control seedlings, but seedlings from the other stages of development no longer had significant differences remaining from the CO2 treatment. This suggests that it could be very efficient to give a short well-timed CO2 pulse at the beginning of the production cycle in hopes of producing a size difference that is maintained throughout the remainder of the greenhouse production cycle under ambient levels of CO2. Short-term exposure to elevated CO2 also increased the ratio of shoot dry weight to total height for the March, April, and May stages of development. The ratio of total nonstructural carbohydrates to free amino acids was negatively correlated (r2 = 0.98) with the allocation of new growth between shoots and roots as measured by the allocation coefficient, k (milligrams shoot growth per milligrams root growth). As seedlings developed along their seasonal growth cycle, ratios of total nonstructural carbohydrates to free amino acids increased and the values for k decreased. The effect of CO2 enrichment on these two factors is discussed. Monitoring total nonstructural carbohydrate and free amino acid concentrations in foliage could have potential as a method to predict the percentage of carbon allocated to root systems of entire forest stands as well as of individual tree seedlings.

The role of sex steroids in the acquisition and production of birdsong.

Male birdsong is generally regarded as a secondary sexual characteristic under the control of gonadal steroids. Song typically waxes and wanes with the seasonal cycle of testicular growth and regression and decreases after adult castration. Testosterone therapy reinstates song, induces it in females, augments it in intact males, and spring testosterone profiles correlate with seasonal song production. Thus, testosterone has been viewed as a major factor in song acquisition and production acting either directly, or after aromatization within the brain. We show here, however, that song learning and early phases of the development of singing both take place in castrated male birds with no significant levels of testosterone in their blood plasma. Testosterone seems to be required for song crystallization, however. Oestradiol was unexpectedly still present after castration, evidently from a non-testicular source, throughout the period of male song acquisition.

Demographic correlates of variable egg survival for a tropical lizard

Anolis limifrons is a small iguanid lizard that is found in lowland rainforest in Central America. Important life attributes include early maturity (3-4 mo), multiple clutches of a single egg per breeding season, and low survival. Population turnover is essentially annual as less than 5% adults live more than one year. My objectives were to resolve the following specific questions: 1) Where are eggs laid? 2) Does survival differ among different types of oviposition site? 3) How is the survival of eggs related to the density and age distribution of lizards at different areas? Most eggs are laid beneath litter on the ground. Eggs laid aboveground are placed in accumulations of soil and litter in stumps, crevices in buttressed trees and vines, holes in trees, and on the tops of logs. Eggs were found in aboveground sites in proportion to the abundance of those sites. The major cause of mortality was predation by Solenopsis ants. Judging by the ratio of eggs known to have hatched to those that were predated, survival did not differ among the various types of oviposition site. Study areas with high egg survival in the middle of the wet season had relatively more young (<4-6 mo old) lizards at the end of the wet season than areas with low egg survival. The strong correlation between the relative number of young individuals and egg survival is a consequence of the seasonal cycle of growth and reproduction. Most eggs are laid in the wet season (May-December). Hatchlings complete growth during the dry season (January-April) and become reproductive adults in their second wet season. However, because hatchlings can grow to reproductive size in approximately 3 months, individuals that hatch early in the wet season may themselves produce offspring that will be <4-6 mo old in the late wet season. The relative survival of juveniles (eggs) and adults is an important datum for many theories of life history evolution. For A. limifrons, survival of eggs was two to three times more variable than survival of lizards on both temporal and spatial scales. Comparative data for reptiles are not available to put this information into an evolutionary prespective.

Food availability controls seasonal cycle of growth in Macoma balthica (L.) in San Francisco Bay, California

A 2-yr field study of growth in the bivalve Macoma balthica (L.) at four locations in San Francisco Bay, California, U.S., showed that the timing and rate of growth (increase in shell length) were related to food supply. This clam feeds on both planktonic and benthic microalgae, depending on availability. Growth was apparently food-limited during some months, during one year more than the other, and at some locations more than others. Tissue-weight changes were also related to food availability. The heaviest animals were found in that year and at those locations with the highest chl a concentrations. Tissue-weight gains usually coincided with increased shell-growth rate or with reproductive development, although some large animals showed weight gain independent of both of these factors during periods with mid-range chl a levels. Weight losses coincided with spawning or periods of low growth rate, except at one station where, during a period when most animals were reproductively ripe, food concentrations were high, and shell growth was rapid, animals lost weight. This study failed to show a relation between salinity and the timing or rate of change of either shell length or tissue weight. The mild temperatures (10–23 °C water temperature) of the area studied resulted in no growth inhibition due to low temperature, but there was some indication that the high air temperatures found in these intertidal areas limited growth rates.

Factors Affecting Photosynthesis, Productivity, and Yield

Abstract The rate and efficiency of photosynthesis (PS) is unquestionably one of the primary determinants of crop yield. Yet, as this symposium illustrates, many other factors contribute to plant productivity. Indeed, the exact relationship between PS rate and yield remains undefined. For several years we have now been interested in factors that affect PS rate in both C3 and C4 plants, not only within the plant, but also external to it. Internal factors affecting PS that will be discussed include leaf age, leaf canopy, photorespiration, stage of plant development (including both vegetative and reproductive phases), and the occurrence of little-recognized metabolic pathways such as polyol synthesis and transport. External factors that affect PS, in addition to obvious environmental determinants such as temperature, light, and water, include the seasonal growth cycle. While we still do not understand how all of these factors interact to affect yield, we are beginning to understand their singular effects. Control of these factors, once thought to be within reach even before the advent of biotechnology, remains likely, but distant.